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JOURNAL.

OF THE

NEW YORK

ENTOMOLOGICAL SOCIETY

Iruotrb to jEttitmtologij in <$*tt*ral

VOLUME LII, 1944

Published Quarterly by the Society
North Queen St. and McGovern Ave. Lancaster, Pa.

New York, N. Y.

THE SCIENCE PRESS PRINTING COMPANY
LANCASTER, PENNSYLVANIA

CONTENTS OF VOLUME LII

Page

Alexander, Charles P.

Records and Descriptions of Neotropical Crane-flies

(Tipulidae, Diptera), XVII 45

Records and Descriptions of Neotropical Crane-flies
(Tipulidae, Diptera), XVIII 369

Arnett, Ross H., Jr,

A Revision of the Nearctic Silphini and Nicrophorini
Based upon the Female Genitalia (Coleoptera, Sil-

phidae) 1

Bacon, Annette L.

Bibliography of Frank E. Lutz 69

Bird, Henry

A Re-Survey of Papaipema Sm. (Lepidoptera) 193

Boyd, William M.

Insect Introductions and War 200

See Weiss, Harry B.

Bromley, Stanley W.

Ephraim Porter Felt 223

Brown, F. Martin

Notes on Mexican Butterflies, II, Pieridae 99

Notes on Mexican Butterflies, III, Danaidae 237

Notes on Mexican Butterflies, IV 343

Caldwell, John S.

PsylliidaB from Tropical and Semitropical America

(Ilomoptera) 335

Clench, Harry K.

Two New Subspecies of Everes Comyntas Godart (Lepi-
doptera, Lycaenidae) 59

Two New Subspecies of Lycaenopsis pseudargiolus Bvd.

& Lee. (Lepidoptera, Lycaenidae) 273

New Neotropical Theclinae (Lepidoptera Lycaenidae) 255

Davis, William T.

The Remarkable Distribution of an American Cicada:

A New Genus and Other Cicada Notes 213

iii

Forbes, William T. M.

Lepidoptera from Western Peru and Ecuador 75

Ginsburg, Joseph M.

Outdoor Protection from Mosquitoes 247

Hemming, Francis

Recent Work by the International Commission on Zo-
ological Nomenclature 211

Huckett, H. C.

A Revision of the North American Genus Eremomy-

ioides Mai loch (Diptera, Muscidae) 361

Huntington, E. Irving

Thecla burdi Kaye, a Synonym 328

Jordan, Karl

The Status and Functions of the International Com-
mission on Zoological Nomenclature and the Present

State of Its Work 385

Linsley, E. Gorton

Hibernation of the Syrphid Fly, Lathyrophthalmus
aeneus Scop. 272

New Species of Neopasites with Notes Concerning Others

(Hymenoptera, Nomadidae) 277

McCoy, E. E.

See Weiss, Harry B.

Members of the New York Entomological Society 389

Michener, Charles D.

Differentiation of Females of Certain Species of Culex

by the Cibarial Armature 263

Miller, Dwight D.

Drosophila melanura, a New Species of the Melanica

Group 85

Milne, Lorus J., and Margery Milne

Notes on the Behavior of Burying Beetles (Nicrophorus

spp.) 311

Milne, Margery

See Milne, Lorus J.

Rapp, William F., Jr.

Catalogue of North American Psychodidse 201

Richards, A. Glenn, Jr.

The Structure of Living Insect Nerves and Nerve
Sheaths as Deduced from the Optical Properties 285

Rupert, Laurence R.

A New Species of Lambdina and Notes on Two Species

of Besma (Lepidoptera, Geometridae, Ennominae) 329

Schneirla, T. C.

The Reproductive Functions of the Army-Ant Queen

as Pace-Makers of the Group Behavior Pattern 153

Weiss, Harry B.

The Death-Feints of Alobates pennsylvanica DeG., and

Alobates barbata Knoch. (Coleoptera) 281

Frank Eugene Lutz 62

Insect Food Habit Ratios of New York State 283

Insect Responses to Colors 267

Notes on the Death-Feint of Bruchus obtectus Say 262

Weiss, Harry B., E. E. McCoy, Jr., and William M. Boyd
Group Motor Responses of Adult and Larval Forms of

Insects to Different Wave-Lengths of Light 27

Wolcott, A. B.

A Generic Review of the Subfamily Phyllobaeinae 121

JnsocJS

Vol. LII No. 1

MARCH, 1944

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Publication Committee

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.

LANCASTER, PA.

NEW YORK, N. Y.

—

1944

CONTENTS

A Revision of the Nearctic Silphini and Nicrophorini
Based upon the Female Genitalia (Coleoptera, Silphidae)

By Ross II . Arnett, Jr 1

Group Motor Responses of Adult and Larval Forms of In-
sects to Different Wave-Lengths of Light

By Harry B. AVeiss, E. E. McCoy, Jr,, and William
M. Boyd 27

Records and Descriptions of Neotropical Crane-Flies
(Tipulidae, Diptera), XVII

By Charles P. Alexander 45

Two New Subspecies of Everes Comyntas Godart (Lep-
idoptera, Lyceenidae)

By Harry K. Clench 59

Frank Eugene Lutz

By Harry B. Weiss 62

Bibliography of Frank E. Lutz

By Annette L. Bacon 69

Lepidoptera from Western Peru and Ecuador

By Wm. T. M. Forbes 75

Drosophila Melanura, a New Species of the Melanica
Group

By Dwight D. Miller 85

Notes on Mexican Butterflies, II, Pieridae

By F. Martin Brown 99

NOTICE: Volume LI, Number 4, of the Journal of The
New York Entomological Society was published on
January 3, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.,
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LII March, 1944 No. 1

A REVISION OF THE NEARTIC SILPHINI AND
NICROPHORINI BASED UPON THE FEMALE
GENITALIA (COLEOPTERA, SILPHID^E)

By Ross H. Arnett, Jr.

Ithaca, New York

The use of the female genitalia as a basis for identification of
genera and species of Coleoptera has been greatly neglected. In
some groups of beetles, however, the female genitalia apparently
offer more diversity of form among species than do those of the
male. Tanner in 1927 pointed out a growing necessity for a
study of the genitalia of beetles for specific descriptions. The
genitalia are naturally more constant within a species than other
parts and they give the taxonomist a better concept of a species,
and its subspecific forms and categories.

The purpose of this study is to present the comparative mor-
phology of the female genitalia of the Neartic Silphini and Nicro-
phorini. It is hoped that the drawings, the key to the genitalia
and the synopsis will be an aid to identification. No basic con-
clusions have been made as to relationships, although certain
tendencies have been indicated.

Acknowledgments. — I wish to express my sincere appreci-
ation and thanks to Professor J. C. Bradley of Cornell University,
under whose direction this work was done, for his many sugges-
tions and criticisms. I am also greatly indebted to Mr. V. S. L.
Pate of Cornell University for help in preparing the paper itself
and to Mr. J. G. Franclemont for aid in developing a technique
for the study of the genitalia.

APR 10 '44

Journal New York Entomological Society

[Vol. LII

I am indebted to Professor M. H. Hatch of the University of
Washington and to the United States National Museum for their
generous loan of material for which I thank them sincerely.

Finally, I wish to thank Mr. E. D. McDonald, Jr. and Mr. Ru-
dolph Schuster for their aid in making the drawings, and to my
wife Mary both for aid in typing the paper and for her continual
encouragement.

Methods. — The methods used in this study are somewhat dif-
ferent from those used by Tanner. The beetle was first relaxed
in hot water and the genitalia removed with a pair of forceps
and boiled in caustic potash. They were then put in water and
the mid-ventral membrane cut, the two lateral plates (the para-
procts) flattened out on each side of the dorsal plate (the proc-
tiger) and the coxite bent out to the side of the valvifer so that
the whole organ was flattened out. It was then mounted in
balsam. This enables projection drawings to be made. It is
essentially the same method as that used in studying the male
genitalia of Lepidoptera.

Sexual Differences. — In the female Silphini, the sutural
angles of the elytra are very acute, whereas in the males they are
generally but not always rounded. The hind femora are greatly
enlarged in the males of Silpha littoralis L. and normal in size in
the female sex. Little sexual differences is evident in Thanatophi-
lus truncata Say.

In the Nicrophorini studied, the males have the eyes situated
well forward on the head, the clypeus large and the fore tarsal
pulvilli expanded, whereas in the females the eyes are placed
well towards the back of the head, the clypeus is appreciably
smaller and the fore tarsal pulvilli are simple.

SPECIES STUDIED

Neartic Species. — A list of the species considered in this paper
follows. Only a few forms of subspecific rank have been studied.
Of those studied, however, some changes have had to be made in
their ranking. Of the others, the opinion of other authors has
been followed in considering their rank. In the case of Nicro-
phorus pulsator Gistel, and Silpha tyrolensis Leach, the evidence
of their existence in the Neartic region is insufficient and they
have not been included here.

Mar., 1944]

Arnett: Silphid.®

NEARTIC SILPHINI AND NICROPHORINI

Silphini

Silpha L.
lift oralis L.

form surinamensis Fab.
ab. bizonatus Port.
—disciocollis Brulle
analis Chev.
cequinoctialis Gistel
braziliensis Dej.
cayennensis Berg, (nec
Sturm. )

var. elongata Port,
var. discreta Port.
Thanatophilus Leach

Subgenus Oiceoptoma Leach
americana L.
peltata Catesby
ab. affinis Kby.
terminata Kby.
canadensis Kby.
acc. brunnipenis Hatch
noveboracensis Forst.
marginalis Fab.
marginata Kby.
quadripunctata L.

quadripunctulata Muller
quadrimaculata Samouelle
var. sexpunctata Gerh.
ab. bifasciata Schulze.
incequalis Fab.

subsp. rugulosa Port.
subrugata Cherv. nom.
nud.

acc. bicolorata Hatch
ramosa Say
cervaria Mann.
cenescens Casey

Sub genus Thanatophilus s. str.
lapponica Hbst.
tuberculata Germ,
subsp. caudata Say
calif ornica Mann,
subsp. granigera Cherv.
tritub erculat a Kby.

sagax Mann.
coloradensis Wick.

obalskii Port.
truncata Say

mexicana Cherv. in litt.

Blitophaga Keitt.
opaca L.
hirta Schaff.
villosa Naezen
tomentosa Villers
var. samnitica Fiori
var. binotata Port.
bituberosa Lee.

Nicrophorini

Nicrophorus Fabricius

Subgenus N ecrochar is Portevin
carolinus Fab.
mediatus Fab.
ab. mysticallis Ang.
ab. scapulatus Port,
ab. dolosus Port,
ab. floridee Hatch
ab. krautwurini Hatch
ab. lunulatus Hatch
ab. nebraskee Hatch

Subgenus Nicrophorus s. str..
orbicollis Say
halli Kby.

quadrisignatus Cast.

Journal New York Entomological Society

[Vol. LII

humator Fab.
sulactus Fisch.
paratype of grandior
Angell

var. atricornis Meier,
ab. maculosns Meier,
ab. rubroplearalis Delah.
delahoni Schilsky i. lift,
ab. signiceps Delab.
subsp. tenuipes Lewis
sayi Cast.

lumulatus Gistel
lunatus Lee.
luniger G. & H.
marginatus Fab.
requiscator Gistel
montezumce Matt,
ab. cordiger Port,
ab. sanjuance Hatch
ab. engelhardti Hatch
ab. leachi Hatch
vespilloides Hbst.
mortuorum Fab.
fractus Port,
ab. andrewesi Port,
ab. aurora Motch.
hebes Kby.
pygmeeus Kby.
vespilloides Lee. (nec.
Hbst.)

defodiens Mann.
disjunctus Wil.-Ellis
ab. humeralis Hatch
ab. Iristis Port,
ab. steinfeldi Smirnov,
ab. maculatus Wil.-Ellis
ab. altumi Westh.
ab. subfasciatus Port.

ab. sub interrupt us Pic.
var. borealis Port,
var. sylvaticus Reitt
ab. sylvivagus Reitt
ab. ruber Hatch
ab. nearticus Hatch
ab. nicolayi Hatch
ab. oregonesis Hatch
subsp. defodiens Mann.
nunemacheri Hatch (nec.

Port.)

ab. binotoides Hatch
binotatus Hatch (nec.

Port.)

ab. conversator Walk.
defodiens var. b. Mann.
lateralis Port.
pollinctor Lee. (nec.

Mann.)

ab. pacificce Hatch
ab. walkeri Hatch
conversator Port. (nec.

Walk.)

ab. gaigei Hatch
ab. kadjakenis Port,
ab. mannerheimi Port,
ab. binotatus Port.
plagiatus Mots.
vespillo L.
vulgaris Fab.
cadaver inus Gravenh.
curvipes Duftschm.
ab. faureli Fauconnet
ab. varendorffi Westh.
ab. bolsmanni Westh.
ab. cethiops Scheicher
ab. minor Westh.
ab. germani Hatch

Mar., 1944]

Arnett: Silphid^e

americanus Oliv.
grandis Fab.
virginicus Frol.
melsheimeri Kby.
hybridus Hatch & Angell
var. minesotianus Hatch
nigritus Mann.

var. ruficornis Mots.
pustulatus Hers.
bicolor Newn.
tardus Mann,
ab. coloradensis Hatch
ab. noveboracensis Hatch
ab. fasciatus Port,
ab. unicolor Port.
investigator Zett.
ruspator Er.
infodiens Mann.
confossor Mots.
micro cephalus Thoms.
pustulatus Horn. (nec.

Hers.)

labiatus Mots.
vestigator Gyll. (nec.

Hers.)

subsp. investigator Zett.
ab. suturalis Mots.
infodiens var. b. Mann,
ab. funeror Reitt.
ab. funerator Fanr.
var. variolosus Port,
ab. intermedins Reitt.
ab. jamezi Hatch
ab. lutescens Port,
subsp. maritimus Guer.
aleuticus Guer.
pollintor Mann.
sibiricus Mots.

infodiens var. c. Mann,
ab. martini Hatch
ab. clarencei Hatch
ab. sitkensis Hatch
ab. massetti Hatch
ab. grahami Hatch
ab. charlottei Hatch
ab. particeps Fisch
ab. japani Hatch
tomentosus Web.
velutinus Fab.
ab. communis Hatch
ab. elongatus Hatch
ab. angustefasciatus Port,
ab. splendens Hatch
ab. brevis Hatch
var. aurigaster Port.
germanicus L.
listerianus Fourer
ab. specious Schultze
ab. bimaculatus Steph.
ab. frontalis Fisch.
ab. fassifer Reitt.
ab. apicalis Kraatz
var. ruthenus Mots.
grandior Ang.
guttula Mots.

subsp. guttula Mots,
ab. ruficornis Mots,
ab. sanfranciscce Hatch
ab. punctatus Hatch
ab. shastce Hatch
ab. hypomerus Hatch
ab. lajollce Hatch
ab. vandykei Ang.
ab. quadriguttatus Ang.
ab. kuschei Hatch

Journal New York Entomological Society

[Vol. LII

mexicanus Matt.
hecate Bland
ab. wallisi Hatch
ab. California Hatch
ab. intermedins Hatch
ab. disjunctus Port,
ab. woodgatei Hatch
ab. phoenix Hatch

ab. novamexicce Hatch
ab. rubripennis Port,
ab. rubrissimus Hatch
ab. immacnlosis Hatch
ob scums Kby.
melsheimeri Lee.
ab. discontinus Hatch
ab. ruber Hatch

Exotic Forms and Their Relationship to Neartic Species. —
The following list of species are exotic forms of which the female
genitalia have been studied. The first name in each group is that
of the type for the genus or subgenus, or it is a typical neartic
species of that group. The next names are those studied with
the generic or subgeneric name as used by other authors follow-
ing it, if it differs from the names employed in this paper.

Silpha littoralis L.

Silpha cayennesis Sturn.
Silpha bigutatta Phil.
Necrodes bigutatta Phil.
Paranecrodes biguttata
Phil.

Thanatophilus ( Oiceoptoma )
americana L.
Thanatophilus thoracica

Silpha thoracica L.
Thanatophilus japonica
Mots.

Silpha japonica Mots.
Thanatophilus obscura L.

Silpha obscura L.
Thanatophilus carinata
Hbst.

Silpha carinata Hbst.
Thanatophilus Icerigata
Fab.

Silpha Icerigata Fab.
Thanatophilus perforata

Gbl.

Silpha perforata Gbl.
Thanatophilus atrata L.

Silpha atrata L.
Thanatophilus nigrita
Creutz

Silpha nigrita Creutz
Thanatophilus granulata
Oliv.

Silpha granulata Oliv.
Thanatophilus ( Thanatophi-
lus) truncata Say
Thanatophilus sinuatus
Fab.

Silpha sinuatus Fab.
Thanatophilus terminata

Hum.

Silpha terminata Hum.
Thanatophilus rugosus L.
Silpha rugosus L.
Blitophaga opaca L.

Blitophaga oblong a Kust.

Silpha oblong a Kust.
Blitophaga souverbiei
Fairm.

Mar., 1944]

Arnett: Silphid^e

Silpha souveriei Fairm.
Blitophaga orientalis
Brnlle

Silpha orientalis Brnlle
Nicrophorus ( Nicrophorus )
vespillo L.

Nicrophorus prcedator

Reitt.

Nicrophorus rotundicollis
Port.

Nicrophorus didymus
Brulle

Nicrophorus interruptus
Steph.

Synopsis of the Neartic Silphini and Nicrophorini. — The
following synopsis is meant to serve as a means of correlating the
external characters with those of the genitalia.

TEIBES

Antennas apparently of ten segments, the second segment being very short
and more or less hidden in the tip of the first; elytra short and not cover-
ing more than the basal five tergitesl Nicrophorini

Antennae clearly of eleven segments, the second not shortened; elytra, if short,
covering more than the basal five tergites Silphini

Genera Silphini

A. Occipital ridge prominent; eyes usually large and prominent; form
usually elongate; labrum broadly emarginate; prothoracic spiracle

sometimes exposed Silpha L.

AA. Occipital ridge usually not prominent; eyes not large and prominent;
labrum broadly or narrowly emarginate; prothoracic spiracle rarely
exposed.

B. Eyes normal, protruding somewhat from the head; labrum broadly
or somewhat narrowly emarginate, but never very narrowly
emarginate unless the head is elongate; head normal or elon-
gate, not short, round or compact Thanatophilus Leach

BB. Eyes very small, not or only very slightly protruding from the
head; head short, round, and compact; labrum very narrowly
emarginate I Blitophaga Beitt.

SILPHA

Pronotum oval, black; elytra usually with red apical spots forming a bar,

sometimes with red basal markings or sometimes immaculate littoralis L.

Pronotum transverse, yellowish-brown with a black central area; elytra im-
maculate discicollis Brulle

THANATOPHILUS

Sub-genera

Labrum broadly or narrowly emarginate; pronotum rarely tomentose, if
tomentose, then orange with a black central area; head and mouth parts
often elongate Oiceoptoma Leach

Journal New York Entomological Society

[Vol. LII

Labrum always very broadly emarginate; pronotum usually tomentose, if
not, then elytra without prominent costse; head and mouth parts never
elongate Thanatophilus s. str.

Sub-genus Oiceoptoma

A. Pronotum orange or yellow with a black central area.

B. Elytra rugose americana L.

BB. Elytra smooth.

C. Costse prominent novaboracensis Frost.

CC. Costae obscure, elytra tan with four black spots and the scu-

tellum black quadripunctata L.

AA. Pronotum black.

D. Elytra smooth incequalis Fab.

DD. Elytra rugose ramosa Say

Sub-genus Thanatophilus

A. Pronotum tomentose; elytral costae prominent.

B. Intervals of the elytral costae tuberculate .lapponica Hbst.

BB. Intervals of the elytral costae flat.

C. Two inner elytral costae subequal throughout.

i tritub erculat a Kby.

CC. Two inner elytral costae nearly obsolete at the base.

coloradensis Wick

AA. Pronotum glabrous, costae obscure ; truncata Say

BLITOPHAGA

Surface pubescent ; form more elongate opaca L.

Surface sparsely pubescent; form more oval bituberosa Lee.

NICROPHORUS

A. Pronotum oboval, without distinct sculpturing and very narrowly mar-
gined (subgenus N ecrocharis) carolinus L.

AA. Pronotum orbicular, transverse or cordate with distinct sculpturing and
widely margined on the sides and back (subgenus Nicrophorus s. str.).

B. Pronotum orbicular, widely margined at the sides and the base.

orbicollis Say

BB. Pronotum not orbicular.

C. Pronotum sinuate at the sides, base nearly as wide as the
apex, sides and base widely margined, not cordate.

D. Metasternal epimeron tomentose.

E. Hind tibia curved.

F. Metatrochanter spine small and divergent;
pronotum disc orange, margin black ; front

orange .:.... americanus Fab.

FF. Metatrochanter spine large and convergent;
pronotum black; front black sayi Lap.

Mar., 1944]

Arnett: Silphid^e

EE. Hind tibia straight.

G. Spine of the metatrochanter obscure; elytra

immaculate nigritis Mann.

GG. Spine of the metatrochanter prominent.

H. Spine divergent; elytra with orange

fascae pustulatus Hersch.

HH. Spine convergent; elytra immaculate.

humator Fab.

DD. Metasternal epimeron glabrous.

I. Elytra immaculate; hind tibia© usually curved.

germanicus L.

II. Elytra with orange fasciae; hind tibia straight.

J. Three terminal segments of the antennae

black vespilloides Hbst.

JJ. Three terminal segments of the antennae
orange.

K. Metasternal pubescence brown; abdom-
inal pubescence black.

mexicanus Matth.
KK. Metasternal pubescence yellow; abdom-
inal pubescence brown.

investigator Zett.

CC. Pronotum with base much narrower than the apex, sides
strongly sinuate, cordate.

L. Metasternal epimeron glabrous melsheimeri Kby.

LL. Metasternal epimeron tomentose.

M. Thorax tomentose.

N. Thorax entirely tomentose.

tomentosus Web.

NN. Thorax tomentose apically only.

vespillo L.

MM. Thorax glabrous.

O. Basal segment of the antennal club black.

P. Hind tibia straight; disc of the pro-

notum punctate guttula Mots.

PP. Hind tibia arcuate; disc of the pro-
notum nearly smooth.

obscurus Kby.

00. Basal segment of the antennal club orange.

Q. Hind tibia arcuate marginatus Fab.

QQ. Hind tibia straight.

R. Ventral surface of the posterior
tibia densely yellow tomen-
tose hecate Bland.

RR. Ventral surface of the hind
tibia sparsely black tomen-
tose hybridus Hatch & Ang.

Journal New York Entomological Society

[Yol. Lll

General Morphology of the Genitalia. — The same terminol-
ogy as that adopted by Tanner, which seems to be a usable inter-
pretation of the relationship of the parts, has been used here.

The dorsal plate or proctiger (PL I, Fig. 3, p.) forms the
upper surface of the genitalia. It has a terminal process (PI.
II, Fig. 8, pro.) sometimes elongate and spatulate, and may
be bent at various angles. Frequently it has terminal hairs. It
apparently serves as the dorsal guide. The paraprocts (PI.
I, Fig. 3, pp.) are lateral plates forming the sides and bottom
of the organ. They sometimes bear setae. The paraproct bears
the valvifer (PI. I, Fig. 3, vf.) which in turn bears the coxite
(PI. I, Fig. 3, c.). The valvifer is sometimes modified into a
lateral guide (PL II, Fig. 8, l.g.) appearing claw-like and lobed
(Pl. II, Fig. 8, 1.) or it may be unmodified and possess setae. The
coxite is a hollow process which supports the stylus (Pl. I, Fig. 3,
sty.) either terminally or laterally on the margin. The stylus is
of various sizes and length and in some species it is expanded at
the apex.

The proctiger is interpreted as a part of the tenth abdominal
tergite. The paraprocts are probably parts of the ninth tergite
with the styli, coxites and valvifers as appendages of the ninth
segment. In this study only the ninth and tenth segments have
been considered. The eighth segment consists of two more or
less unmodified plates, the tergite and sternite.

Key to the Neartic Silphini and Nicrophorini Based
on the Female Genitalia

1. Valvifer at most only slightly lobed, not developed into a curved proc-

ess; proctiger never extended and lobed, ( Silphini ) (2).

Valvifer with a well developed curved process and the proctiger usually
extended and lobed. ( Nicrophorini one genus Nicrophorus Fab.)

(14).

2. Stylus apical or lateral; coxite without a lateral projection (3).

Stylus lateral; coxite with a lateral projection. ( Blitophaga Beitt.)

(12).

3. Stylus always apical, stout, the diameter nearly that of the coxite; cox-

ite stout and more or less uniform throughout. ( Silpha L.) (4).

Stylus apical or lateral, if apical, then very small, much smaller than
the coxite and the coxite is much wider at the base than at the apex.
( Thanatophilus Leach.) . (5).

Mar., 1944]

Arnett: Silphid^

SILPEA

4. Coxite with two lateral connecting ridges; stylus broader at the apex.

littoralis L.

Coxite without ridges; stylus more uniform throughout.

discicollis Brulle

THANATOPHILUS

5. Stylus apical or lateral, if lateral then the coxite beyond the stylus is

not greatly flattened or lobed. Subgenus Oiceoptoma Leach (6).

Stylus always lateral; coxite beyond the stylus greatly flattened and
slightly lobed. Subgenus Thanatophilus s. str. (10).

6. Stylus apical 4-punctata L.

Stylus lateral , (7).

7. Stylus long and angulate, nearly reaching the apical end of the coxite.

americana L.

Stylus short and not angulate, and much shorter than the portion of
the coxite beyond the insertion of the stylus (8).

8. Apical portion of the coxite beyond the base of the stylus twice the

length of the stylus or less (9).

Apical portion of the coxite beyond the base of the stylus much more
than twice the length of the stylus ramosa Say

9. Bridge (PL I, Pig. 3, br.) between the coxite and the valvifer wide;

proctiger broadly rounded apically; stylus round at the apex.

novdboracensis Forst.

Bridge between the coxite and the proctiger narrow; proctiger more

angular apically; stylus angulate at the apex inaequalis Fab.

L0. Stylus greatly enlarged at ‘the apex, more than twice the width of the

basal portion truncata Say.

Stylus enlarged at the apex, but much less than twice the width of the
basal portion (11) .

11. Coxite with a prominent basal spine below the insertion of the stylus.

coloradensis Wick.

Coxite without a prominent basal spine below the insertion of the
stylus (12).

12. Stylus inserted on the ventral surface of the coxite so that there is

apparently a lateral flap covering the base of the stylus.

lapponica Hbst.

Stylus inserted on the lateral surface of the coxite tritub erculat a Kby.

BLITOPHAGA

13. Stylus as long as the lateral lobe of the coxite bituberosa Lee.

Stylus much shorter than the lateral lobe of the coxite opaca L.

NICROPHORUS

14. Coxite with a terminal claw as long or longer than the stylus. (Sub-

genus Neocrocharis Port.) carolinus L.

Journal New York Entomological Society

[Vol. Lir

Coxite without a terminal claw, or if apparently present, then always
much less than the length of the stylus. (Subgenus Nicrophorus s.
str.) (15).

15. Proctiger lobe short and broad (16).

Proctiger lobe long and narrow, without an apical spatula.

marginatus Fab.

Proctiger lobe medium in length and width, with or without an apical
spatula (18).

16. Proctiger lobe bifurcate orbicollis Say

Proctiger lobe not bifurcate : (17).

17. Proctiger lobe truncate, without prominent apical ridge humator Oliv.

Proctiger lobe round, with prominent apical ridge. (PI. II, Fig.

8, r.) ...... sayi Lap.

18. Proctiger without an apical spatula vespilloides Hbst.

Proctiger with an apical spatula (19).

19. Coxite with a basal-lateral lobe; lobe of the claw of the valvifer longer

than wide vespillo L.

Coxite without a basal-lateral lobe; lobe of the claw of the valvifer
always broader than long (20).

20. Coxite emarginate on the inner lateral margin americanus L.

Coxite not emarginate \ (21).

21. Lobe of the proctiger sub-truncate (22).

Lobe of the proctiger round (24).

22. Lobe of the proctiger greatly curved dorsally-ventrally (23).

Lobe of the proctiger slightly curved dorsally-ventrally nigritus Mann.

23. Spatula of the proctiger lobe broad melsheimeri Kby.

Spatula of the proctiger lobe narrow .• hybridus Hatch & Ang.

24. Lobe of the valvifer claw obscure . ......... (25).

Lobe of the valvifer claw prominent (27).

25. Coxite very narrow hecate Bland.

Coxite broad , (26).

26. Proctiger lobe greatly curved dorsally-ventrally (28).

Proctiger lobe slightly curved dorsally-ventrally (29).

27. Lobe of the claw of the valvifer with setae, spatula ridged.

germanicus L.

Lobe of the claw of the valvifer without setae, spatula not ridged.

pustulatus Hers.

28. Apical margin of the base of the valvifer concave and nearly parallel

with the basal margin mexicanus Matt.

Apical margin of the base of the valvifer nearly straight and not paral-
lel with the basal margin tomentosus Web.

29. Spatula of the proctiger lobe oval guttula Mots

Spatula of the proctiger lobe round (80).

30. Valvifer, exclusive of the lobe, square obscurus Kby.

Valvifer, exclusive of the lobe, trapizoidal investigator Zett.

Mar., 1944]

Arnett: Silphid^e

DISCUSSION OF THE NEARTIC SILPHINI AND NICROPHORINI

General. — The tendencies pointed out here are based only on
the species considered in this study and without comparison with
other beetles.

On the basis of the female genitalia alone, the Silphini appear
to be the more primitive of the two tribes and have been treated
so here. But this conclusion is based on the assumption that
simplicity of form, as seen in the Silphini, indicates primitive-
ness rather than reduction. The Nicrophorini are so closely
linked together that it is difficult to tell anything about their
phylogeny.

Silphini

The proctiger is simple, never lobed, usually with setae; the
paraproct is simple, essentially the same as that of the Nicro-
phorini, but with setae. The valvifer at most is lobed only,
usually with setae; the coxite exhibits the greatest variation of
the organ. It may have a basal lobe, or be uniform throughout.
The stylus is attached to the coxite either terminally or laterally.
The stylus is usually uniform in shape, but in some groups it is
enlarged at the apex.

Silpha L.

Type : Silpha littoralis L., designated by Latreille 1810.

In this genus the stylus is terminal and stout, and is wider at
the apex than at the base. The coxite is stout and uniform
throughout. The proctiger, paraprocts and the valvifers are
unmodified.

Silpha littoralis L.

This species is supposedly European, but the characters used
to separate it from the Neartic surinamensis Fab. do not ade-
quately separate the two. In the collection of the author there
are specimens with immaculate elytra, and the genitalia of the
two forms show no differences. Therefore surinamensis Fab. is
a form of littoralis L. and not a distinct species. The genitalia
of this species has a setigerous proctiger. The coxite is stout and
with two ridges on the outer lateral surface which connect at
the apex.

Silpha discicollis Brulle.

Journal New York Entomological Society

[Vol. LII

Proctiger with setae; coxite without lateral ridges; stylus less
enlarged at the apex.

Thanatophilus Leach

Type : Silpha rugosa L.

Coxite blade-like, flattened or uniformly triangular; stylus
terminal or lateral, uniform throughout or enlarged at the apex.

Subgenus Oiceoptoma Leach

Type : Silpha thoracica L.

Coxite uniformly triangular with the stylus terminal, varying
to coxite slightly flattened apically and the stylus lateral ; stylus
uniform throughout. Species as described in the key and
synopsis.

Subgenus Thanatophilus s. str.

Coxite flattened at the apex, appearing blade-like and strongly
curved on the outer side ; stylus lateral and enlarged at the apex.
The characters of the species are as presented in the key and
synopsis.

Blitophaga Eeitt.

Type : Silpha opaca L.

Coxite with basal lobe or tooth, terminal portion narrow and
flattened; stylus small and lateral between the basal lobe and
the apex of the coxite, never longer than the basal lobe. The
characters of the species are as presented in the key and the
synopsis.

Nicrophorini

The greatest difference between this tribe and the Silphini is
in the modification of the proctiger. Here the proctiger is
usually greatly extended and generally spatulate at the apex,
nearly always with setae. Also, the valvifer is quite different in
appearance from that of the Silphini. It is greatly enlarged
and extended, flattened and claw-shaped. The coxite is uniform,
bearing the stylus terminally in all cases except Nicrophorus
carolinus L. which has a lateral stylus. The paraproct is with-
out setae, but at times is ridged. The species of this genus are
very closely related with the exception of Nicrophorus carolinus

Mar., 1944]

Arnett: Silphid.®

L. which shows characters differing from the others and is placed
in the subgenus Necrocharis Port.

Nicrophorus Fab.

Characters the same as those of the tribe. Type : Nicrophorus
vespillo L., designated by Latreille, 1810.

Subgenus Necrocharis Port.

Type : Nicrophorus Carolina L., one species only with charac-
ters as in the key and the synopsis.

Subgenus Nicrophorus s. str.

The species of the subgenus are all so closely related that they
cannot be separated into species groups. The characters used
in describing the species are inadequate. Color pattern has
little or no value in separating the majority of the species.
Their relationships depend entirely on what set of characters
are used. Many aberrations have been described, but this is
quite unnecessary and becomes extremely confusing, especially
when they are not illustrated. Because of the great variation in
the color pattern, almost any population can be described as a
new aberration. Some changes have been made in the status of
certain forms. Undoubtedly, when other forms are examined,
more changes will be necessary. The following changes have
been made on the basis of the characters presented in the key and
synopsis :

Nicrophorus melsheimeri Kby. is a distinct species and not a
synonym of investigator Zett.

Nicrophorus nigritus Mann, is a distinct species and not a sub-
species of investigator Zett.

Nicrophorus hecate Bland, is a distinct species and not a sub-
species of guttula Mots.

CONCLUSIONS

1. The female genitalia of Silphini and Nicrophorini present
characters which serve to separate the species of the groups.

2. The two tribes have basically the same type of female geni-
talia, but they are two very distinct groups of genera.

3. Silphini tends to be more primitive than Nicrophorini.

Journal New York Entomological Society

[VOL. LII

4. The genera Silpha, Thanatophilus , Blitophaga and Nicro-
phorus are distinct groups of species.

5. The color patterns of Nicrophorus are not good specific
characters and aberrations based on those characters are worth-
less.

6. The form and sculpturing of the elytra of Silphini do not
show relationships between the species.

7. Nicrophorus offers no distinct species groups, and the spe-
cies of the genus are very closely related.

BIBLIOGRAPHY

Hatch, M. H. Studies on the Silphinae. Jour. N. Y. Ent. Soc., XXXV,
331-370, 1927. Coleopterorum Catalogus, Part 95, Junk, Berlin,
1928.

Hatch and Reuter. Coleoptera of Washington, Silphidae. Univ. of Wash.
Publ. in Biol., 1: 147-162, 1934.

Horn, G. H. Synopsis of the Silphidae of the U. S. with reference to the
genera of other countries. Trans. Am. Ent. Soc., 8: 219-322, pi.
V-VII, 1880.

Latreille, P. H. Considerations Generales sur Pordre Naturel des Ani-
maux. Paris, 1810.

Leng, C. W. Catalogue of the Coleoptera of America, North of Mexico and
suppl. 1-4. Sherman, Mount Vernon, N. Y., 1920.
Scegoleva-Barovshaja, T. Les Necrophorini (Coleoptera, Silphidae) de la
faune de PU.R.S.S. Travaux de l’lnstitut Zool. de PAcademei des
sciences de PU.R.S.S., 1: 161-191, 1932.

Semenov-Tian Shanskij, A. P. De Tribu Necrophorini (Coleoptera, Sil-
phidae) classification et de ejus distributione geographica. Travaux
de l’lnst. Zool. de PAcademie des Science de PU.R.S.S., 1: 149-
160, 1932.

Tanner, V. M. A Preliminary Study of the Genitalia of Female Coleop-
tera. Trans. Am. Ent. Soc., 53, 5-50, 1927.

Mar., 1944]

Arnett: Silphid^e

Abbreviations on Plates

proctiger

spa

spatula

paraprocts

lobe

sty

stylus

ridge

coxite

br. ...

bridge

valvifer

pro

process

lateral guide

Journal New York Entomological Society

[Yol. LI i

PLATE I

Figure la. Silpha L. Elytra of female.

Figure lb. Silpha L. Elytra of male.

Figure 2a. Nicrophorus Fab. Head of male.
Figure 2b. Nicrophorus Fab. Head of female.
Figure 3. Thanatophilus quadripunctata L.
Figure 4. Silpha littoralis L.

Figure 5. Silpha discicollis Brulle.

Figure 6. Thanatophilus americana L.

Figure 7. Thanatophilus novaboracensis Forst.
Figure 8. Thanatophilus incequalis Fab.

Figure 9. Blitophaga bituberosa Lee.

Figure 10. Thanatophilus tritub erculata Kby.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate I)

Journal New York Entomological Society

[Yol. Lll

Figure

PLATE II

1. Thanatophilus ramosa Say.

2. Thanatophilus lapponica Hbst.

3. Thanatophilus coloradensis Wick.

4. Thanatophilus truncata Say.

5. Blitophaga opaca L.

6. Thanatophilus thoracica L.

7. Thanatophilus rugosus L.

8. Nicrophorus carolinus L.

9. Nicrophorus orhicollis Say.

10. Nicrophorus vespilloides Hbst.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate II)

Journal New York Entomological Society

[Vol. LII

Figure 1.
Figure 2.
Figure 3.
Figure 4.
Figure 5.
Figure 6.
Figure 7.
Figure 8.

PLATE III

Nicrophorus americana Fab.
Nicrophorus sayi Fab.
Nicrophorus nigritus Mann.
Nicrophorus pustulatus Hersch.
Nicrophorus humator Fab.
Nicrophorus germanicus L.
Nicrophorus mexicanus Matth.
Nicrophorus investigator Zett.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate III)

Journal New York Entomological Society

[Vol. LII

Figure 1,
Figure 2,
Figure 3,
Figure 4,
Figure 5.
Figure 7
Figure 6.
Figure 8,

PLATE IY

Nicrophorus melsheimeri Kby.
Nicrophorus tomentosus Web.
Nicrophorus vespillo L.

Nicrophorus guttula Mots.
Nicrophorus ohscurus Kby.
Nicrophorus hecate Bland.
Nicrophorus marginatus Fab.
Nicrophorus hyhridus Hatch & Ang.

(Plate IY)

(Jour. N. Y. Ent. Soc.), Vol. LII

1 2

Mar., 1944]

Weiss et al.: Insect Behavior

GROUP MOTOR RESPONSES OF ADULT AND
LARVAL FORMS OF INSECTS TO DIF-
FERENT WAVE-LENGTHS
OF LIGHT

By Harry B. Weiss, E. E. McCoy, Jr., and
William M. Boyd

This, the fifth paper of a series, relating to the group behavior
of insects to colors, is concerned with the responses of seven
species of Coleoptera and of sixteen species of lepidopterous,
hymenopterous and coleopterous larvae to ten wave-length bands
of light of equal physical intensities, in disarray, from 3650 A
to 7400A. The tests were run in the sector type equipment de-
scribed in the third and fourth papers of the series.1 As outlined
in these papers, the insects were placed in an introduction cham-
ber, six feet away from the filter chambers, after the lamps were
on and after all filter chambers were open. After the exposure
period, the filter chambers, the central compartment, introduction
chamber and dark chamber were closed, and counts were then
made.

In all previous tests reported upon in the third and fourth
papers the color filters were arranged in a sequence beginning
with the shorter wave-lengths and extending successively to the
longer wave-lengths, as follows: 3650 A (ultra-violet); 4360 A
(violet-blue) ; 4640 A (blue) ; 4920 A (blue-blue-green) ; 5150 A
(blue-green) ; 5460 A (yellow-green) ; 5750 A (yellow-yellow-
green) ; 6060 A (yellow-orange) ; 6420 A (orange-red) ; and
7200 A (infra-red). Each filter chamber was separated from its
neighbor by a black chamber.

All tests reported in the present paper, with adults and larvae,
were made with the filters in disarray, as follows: 3650 A (ultra-
violet) ; 6060 A (yellow-orange) ; 4640 A (blue) ; 7200 A (infra-
red) ; 5150 A (blue-green) ; 6420 A (orange-red) ; 5750 A (yellow-
yellow-green) ; 4360 A (violet-blue) ; 5460 A (yellow-green) ;
and 4920 A (blue-blue-green). The wave-length figures repre-
sent the peak transmissions of the filters.

i Jour. N. Y. Ent. Soc., 50(1) : 1-35, 1942; 51(2) : 117-131, 1943.

Journal New York Entomological Society

[Vol. LII

Owing to the deterioration that occurred in the lamps used in
previous tests, new forty-watt, frosted, Westinghouse Mazda
lamps and a new General Electric Mazda mercury lamp (type
A-H4, 100 watts) were utilized for all tests. The same method,
outlined in our first paper,2 was used for determining the relative
positions of the lamps and various filter combinations so that the
physical intensities were approximately equal. A slight change
in technique, designed to improve the equalization, was recently
made and this resulted in a new set of distance settings, differing
slightly from those given in the first paper. These new distance
settings are shown on page 29.

RESULTS WITH ADULT INSECTS

Table I presents the results of exposing seven species of Cole-
optera to ten wave-length bands of light of equal physical intensi-
ties, in disarray, in the sector type equipment. By consulting
the percentage distribution of those reacting positively to the
various wave-length bands and by an examination of the group
behavior curves in Figures 1 and 2, it may be noted that the peak
response for all species except Popillia japonica took place in
the ultra-violet (3650-3663 A) and that secondary peaks occurred
either in the blue-blue-green (4920 A) or in the blue-green
(5150 A). Smaller numbers, in general, appear to have gone to
4360 A (violet-blue) and larger numbers to 6060 A (yellow-
orange) than in previous tests when the filters were not in disar-
ray. Except for the somewhat reduced attractiveness of 4360 A
and the slightly increased attractiveness of 6060 A, the behavior
patterns, with the filters in disarray, did not differ materially
from previous patterns obtained with the filters in orderly array.

Peterson and Haeussler3 in their work with the Oriental fruit
moth and colored lights found that when a less attractive colored
light was placed at right angles to a more attractive colored one,
more fruit moths went to the less attractive light than when the
less attractive one was opposite the more attractive one. There is
no doubt that the stimulating power of certain wave-lengths is
influenced by their positions with respect to other wave-lengths.
But the fact remains that except for the slight differences noted

2 Jour. N. Y. Ent. Soc., 49(1) : 1-20, 1941.

3 Ann. Ent. Soc. Amer., 21(3) : 353-379, 1928.

Mar.,

1944]

Weiss et al.:

Insect Behavior

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Mar., 1944]

Weiss et al.: Insect Behavior

above the group behavior patterns for the species tested remained
materially unchanged with the filters in disarray. Regardless of
the relative positions of the various wave-length bands, the insects
made approximately the same selections time after time.

RESULTS WITH LARVAL FORMS
Most of the experimental work on the behavior of insects to
colored light has been done with adult insects. Nevertheless
there are a few references in the literature to the behavior of
larval forms and mention will be made of the photopositive ones.

For example, Mayer and Soule found that the larvae of Danais
archippus are photopositive to ultra-violet. Gross,4 in his study
of the reactions of arthropods to monochromatic lights of equal
intensities reported that the larvae of Zeuzera pyrina, a lepidop-
terous wood-borer, and of a noctuid moth Feltia subgothica, are
photopositive to colors, the order of the effectiveness of stimula-
tion being blue (4200-4800 A), green (4900-5500 A), yellow
(5700-622 A), and red (6300-6500 A). Lymantria larvae, ac-
cording to Hundertmark5 appear to prefer blue when different
colors are compared. Gotz,6 in his study of the perception of
color and form in lepidopterous larvae found that an appreciation
of color occurs in the larvae of Vanessa and Pieris. These are
attracted by the green color of leaves or pieces of paper, regard-
less of the color of the surroundings, but more so on a white back-
ground than on a black one. Lammert7 reports that caterpillars
will go toward a source of light after a blacking of their eyes.
And Suffert8 states that many caterpillars colored like their sur-
roundings and feeding in exposed situations, orient themselves so
that the light always falls upon them from a particular angle.
These last two instances indicate the possession of a dermal light
sense.

Our tests as reported in the present paper involved the ex-
posure of the larvae of sixteen species of insects to ten wave-length
bands of light, of equal physical intensities, from 3600 A to 7200
A. These bands were in disarray. From Table II and Figures
4 Jour. Exp. Zool., 14: 467-512, 1913.
s Z. vergl. Physiol., 24 : 563-582, 1936.
e Z. vergl. Physiol., 23 : 429-503, 1936.

7 Z. vergl. Physiol., 3 : 225-278, 1925.

8Z. Morph. Oekol. Tiere., 26: 147-316, 1932.

TABLE II

Behavior of Sixteen Species of Larvje to Ten Wave-length Bands of Light, or Colors, in Disarray

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Journal New York Entomological Society

[Vol. Lll

3, 4 and 5 which record their group behavior in percentages and
graphically, it is apparent that the peak response for most species
took place in the ultra-violet (3650 A). In nearly all instances
a peak either equal to the one in ultra-violet or secondary to it
occurred in the blue-blue-green (4900 A), or in the blue-green
(5150 A). The minor peak at 6060 A (yellow-orange) is attrib-
uted to the disarray of the filters which resulted in this wave-
length being next to 3650 A (ultra-violet).

The larvae of Diacrisia virginica were the only ones which ultra-
violet light (3650 A) failed to stimulate appreciably. These
larvae are found crawling upon the ground or feeding upon low
plants. As a whole the group behavior of the larvae, in general,
did not differ from that of numerous adult insects, previously
tested, and their color discrimination, so called, was approxi-
mately the same as that exhibited by adult insects in spite of the
fact that their visual organs are less complex than those of adult
insects.

NOTES

Autoserica castanea. This beetle, being nocturnal, was tested
at 10 : 30 p.m.

Hippodamia convergens Guer. The predaceous larvae of this
coccinellid failed to react at all under the conditions of our tests.
When placed in the introduction chamber they climbed up the
sides and remained there. Apparently their negative geotropic
behavior predominated.

Hyphantria cunea Dru. The \ to f grown larvae of this
species, the fall webworm, made a web in the introduction cham-
ber and stayed there, even though they had been previously
deprived of food for twenty hours.

During the course of our work with larvae it was found that,
as a rule, they were more photopositive after having been de-
prived of food for a half-day or more previous to the tests. The
gregariousness of some of the species, especially of the larvae of
Hadena turbidenta Hbn., and Melalopha inclusa Hbn., appeared
to inhibit somewhat their sensitivity to light.

DISCUSSION

In view of the comparative simplicity of the lateral ocelli of
larvae, the similarity of the group behavior of larvae to that of

Mar., 1944]

Weiss et al.: Insect Behavior

adult insects with compound eyes is of considerable interest.
Although variable in structure, lateral ocelli in lepidopterous
larvas consist of a group, each ocellus having a structure not un-
like the single ommatidium of a compound eye. In the larvse of
sawflies and of many Coleoptera, the ocellus, of which there is
only one on each side, is a lens-like, transparent thickening of the
cuticle with underlying epidermis, and retinulse, each made up
of two or three visual cells grouped around a rhabdom. These
visual cells may be pigmented, or there may be separate pigment
cells. Dethier in a recent study9 of the corneal lens in caterpil-
lars states that “the cornea possesses a short focal distance, great
depth of focus, and an extremely low f value permitting the
admittance of much light. ’ ’

Although the king-crab, Limulus polyphemus, is not an insect,
the work of Hartline and Graham on the nerve impulses and
responses of single visual sense cells, to light, in the eye of this
animal is of unusual interest and it is within the realm of possi-
bility that a similar process of photoreception may operate in
insects.

The lateral facetted eye of the king-crab contains about 300
large ommatidia and the optic nerve fibres come directly from the
receptor cells with no intervening neurones. These authors10
studied the nerve impulses and developed a technique by which
was recorded the discharge from a single receptor unit, in the
form of oscillograms, representing the potential changes between
the cut end and an uninjured portion of the nerve, upon stimu-
lation of the eye by light. The electrical activity in the optic
nerve brought about by this stimulation was amplified by a vac-
uum tube and recorded by an oscillograph. Among other things
the stimulation of a single ommatidium resulted in a small strand
of the optic nerve showing a regular sequence of nerve impulses.
‘ 4 The discharge in a single fiber begins after a short latent period
at a high frequency, which has been found to be as high as 130
per second. The frequency falls rapidly at first, and finally
approaches a steady value, which is maintained for the duration
of illumination” (Hartline and Graham).

9 Jour. Cell, and Comp. Physiol., 19(3) : 301-313, 1942.

10 Jour. Cell, and Comp. Physiol., 1(2) : 277-295, 1932.

Journal New York Entomological Society

[Vol. LII

In a later paper11 these authors studied the responses of single
visual sense cells to visible light of different wave-lengths. This
was done by means of single fiber preparations from a Limulus
eye. It was found that when the energy of the stimulating light
of different wave-lengths was approximately equal, the response
to green was stronger than the responses to either violet or red.
When the energy was increased in the red and violet their level
of response was raised and when the intensities for the different
wave-lengths were adjusted so that the responses were equal,
there was no effect of wave-length as such, indicating that single
sense cells can gauge brightness but cannot distinguish wave-
length. The relative energies of the various wave-lengths re-
quired to produce the same response after being adjusted in in-
verse ratio to the degree to which they are absorbed yielded a
visibility curve, for a single visual sense cell, that had its maxi-
mum in the green near 5200 A and that declined symmetrically
on each side to low values in the violet near 4400 A and in the red
near 6400 A. According to the interpretation of visibility curves
by Hecht and Williams12 the stimulation of a single visual sense
cell by light depends upon the absorption spectrum of the pri-
mary photosensitive substance. The absorption of light by this
substance varies with wave-length and the production of a given
response needs a certain amount of photochemical change, which
in turn requires the absorption of a constant amount of energy.

Hartline and Graham also found that in the same eye of Limu-
lus there was a differential sensitivity among optic nerve fibers
and their attached sensory cells for different regions of the visible
spectrum and they believe that such specialization of the visual
cells, coupled with integrated action may give rise to color vision.

In considering the tests with insects reported upon in this and
in previous papers,13 the following patterns of behavior prevailed
over and over, when various species were exposed to ten wave-
length bands of equal intensities from 3600 A to 7200 A. In the
composite behavior14 of 5,454 insects of various orders, mostly

11 Jour. Gen. Physiol., 18(6) : 917-931, 1935.

12 Jour. Gen. Physiol., 5: 1, 1922.

13 JOUR. N. Y. Ent. Soc., 49 : 1-20, 149-159, 1941 ; 50 : 1-35, 1942 ; 51 :
117-131, 1943.

ii Ent. News, 54: 152-156, 1943.

Mar., 1944]

Weiss et al. : Insect Behavior

coleopterous, the peak response took place at 3650 A (ultra-
violet). From here the response declined gradually to a low
point at 4640 A (blue) ; then it increased to a secondary peak
at 4920 A (blue-blue-green), and then declined gradually to a
low point at 5750 A (yellow-yellow-green) from which point it
levelled off to 6420 A (orange-red). In the cases of individual
species there were deviations from this pattern. Drosophila and
various species of Coleoptera in some tests responded almost en-
tirely to 3650 A alone, dropping to a low level at 4360 A and
levelling off at that wave-length. Sometimes the secondary peak
occurred at 5150 A instead of at 4920 A. Although the peak
responses took place at 3650 A and 4920 A, small percentages of
the test animals went to other wave-lengths. In addition, it was
found that when a second test, using the same insects, succeeded
the first, the same group behavior pattern took place. The peak
response occurred at 3650 A, the secondary one at 4920 A. How-
ever, the individuals that made up the peaks in the second test
were not all the same as those making up the peaks in the first
test. In other words, there was a shifting of the individuals that
went to the different wave-length bands, but no difference in the
final result.

Assuming that the light receptors of insects function in the
same way as those of some other invertebrates and considering
the results obtained from the single visual sense cells of Limulus
as outlined by Hartline and Graham and mentioned above, it is
possible to venture an explanation for the group behavior pattern
of insect response to colors. Starting with the fact that the test
insects responded in varying numbers to all wave-lengths from
3650 A to about 6420 A, it is apparent that the photosensitive
substance of their visual sense cells will function at any of the
wave-lengths between 3650 A and 6420 A, if the physical intensity
of the wave-length is sufficient and constant.

When confronted by ten wave-length bands of equalized inten-
sities which converged upon the insects in the introduction cham-
ber of the apparatus, the primary photosensitive substance of the
visual sense cells of a large number absorbed the energy at 3650 A
to a greater extent than the energy at other wave-lengths. This
resulted in a photochemical reaction accompanied by physical

Journal New York Entomological Society

[Vol. LII

changes in nerve fibers, one of which was a change in the electric
potential of the point in the fiber that was actively responding
measured with respect to a nearby, but as yet, inactive point.
This electrical activity in the optic nerve fiber was transmitted
to the muscles where changes in tension occurred resulting in the
insects going to the ultra-violet (3650 A) in larger numbers than
to any other test wave-length. As the absorption of light by the
primary photosensitive substance of the single sense cell varies
with wave-length and as the production of a response requires a
certain amount of photochemical change plus a constant amount
of energy, it seems evident that the energy of the remaining test
wave-lengths although equal, was not sufficient to result in a
response that equalled that of the ultra-violet. Consequently
smaller numbers of individuals responded to the test wave-lengths
other than 3650 A. The question then arises as to why all indi-
viduals did not respond to 3650 A alone. In a group of 100 or
more insects collected in the field, it is not expected that they
would all be in the same physiological state at the same time. In
fact, when tested, only some are photosensitive. Others remain
in the introduction chamber, others get as far as the central com-
partment and others go to the black chamber, all exhibiting dif-
ferent degrees of behavior to light. Among those that are photo-
positive it is reasonable to assume that there exist some varia-
tions by individuals in the sensitivity of their visual receptors.
These variations may be connected with different physiological
states. They may be due to a depletion of the primary photosen-
sitive substance in the visual sense cells through the action of
light, resulting in individuals so affected responding in smaller
numbers to wave-lengths other than ultra-violet. Until restora-
tive processes take place in the visual sense cells of such individ-
uals, their sensitivity to ultra-violet declines.

Frequently various species, when tested, responded almost ex-
clusively to ultra-violet. But many others did not. In the case
of Drosophila which was bred under controlled conditions and
which were of uniform ages, the response to ultra-violet was
unusually high.

It is realized that these deductions are based upon the behavior
of single visual sense cells of the king-crab, to light of different

Mar., 1944]

Weiss et al.: Insect Behavior

wave-lengths and no consideration has been given to the fact that
individual cellular units act collectively and not independently,
nor to the fact that Graham and Hartline15 found that although
the visibility curves for single sense cells in the same eye are
approximately identical, they differ by significant amounts. In
addition they report that two sense cells were able to distinguish
violet from red and taking all these facts into consideration they
are of the opinion that such differential sensitivity “may be con-
sidered a peripheral mechanism of color vision.”

Another reason for the deductions as outlined consists of the
behavior of the Japanese beetle, Popillia japonica, which was
made to respond to what were unattractive wave-lengths under
equalized physical intensities, by increasing the intensities of such
wave-lengths. In fact with other species as well it was possible
to vary the behavior pattern by changing the intensities. And
in general, from our work over the past several years, it appears
that the behavior patterns of insects to equalized wave-lengths
are not unlike the behavior pattern of a single sense cell, in Limu-
lus, to equalized wave-lengths. Perhaps the behavior curves in
this and in former papers16 may be interpreted as rough approxi-
mations of the absorption spectrum of the photosensitive sub-
stance in the combined visual sense cells of many insects, as well
as indications of their motor responses to equalized wave-lengths
of light.

A word should be said about the comparatively large per-
centages of test insects which remain in the introduction chamber
and central compartment of our testing equipment. We have
always attributed this mainly to low illumination. At low illu-
minations only the most sensitive ommatidia function, there being
different thresholds of response for different ommatidia.17 By
others, a falling off in intensity discrimination, due to low illu-
mination is attributed to a nervous coupling of groups of omma-
tidia to form new units.18

is Jour. Gen. Physiol., 18: 917-931, 1935.

is Jour. N. Y. Ent. Soc., 49 : 1-20, 149-159, 1941 ; 50 : 1-35, 1942 ; 51 :
117-131, 1943.

i7 Hecht and Wald. Jour. Gen. Physiol., 17 : 517-547, 1934.
is Buddenbrock and Shultz. Zool. Jahrb. Physiol., 52: 513-536, 1933.

Journal New York Entomological Society

[Vol. Lll

The foregoing discussion is an attempt to explain the group
behavior patterns, or motor responses of insects to various wave-
lengths of light of equal physical intensities on the basis of the
results obtained by investigators who used single visual sense cells
of other invertebrates. It is realized that the motor response to
light of a complex organism such as an insect cannot be ade-
quately and definitely explained on the basis of the behavior of
single visual sense cells of other animals, nevertheless such work
as has been done with single sense cells furnishes valuable clues
to the phenomena of vision in insects. Until similar and addi-
tional investigations are made on the behavior of photoreceptor
cells and optic nerve fibers of insects, singly and in integrated
action, one has to be satisfied with implications.

PLATE Y

Figure 1. Behavior of six lots of Leptinotarsa decemlineata Say, to 10 wave-
length bands, in disarray, from 3650 a to 7200 A. Physical
intensities equalized.

1. 129 beetles. Three tests.

2. 478 beetles. Four tests.

3. 857 beetles. Six tests.

4. 857 beetles. Four tests.

5. 340 beetles. Four tests.

6. 338 beetles. Four tests.

Figure 2. Behavior of six species of Coleoptera to 10 wave-length bands, in
disarray, from 3650A to 7200 A. Physical intensities equalized.

1. Chrysochus auratus Say. 217 adults. Three tests.

2. Tetraopes tetraophthalmus Forst. 144 adults. Five tests. •

3. Chauliognathus marginatus Fabr. 42 adults. Two tests.

4. Photinus pennsylvanica DeG. 78 adults. Three tests.

5. Popillia japonicg Newm. 295 adults. Three tests.

6. Autoserica castanea Arrow. 338 adults. Three tests.

Tested after 10: 30 p.m.

(Jour. N. Y. Ent. Soc.), Vol. LII (Plate V)

Journal New York Entomological Society

[Vol. LII

PLATE VI

Figure 3. Behavior of coleopterous, hymenopterous and lepidopterous larvse
to 10 wave-length bands, in disarray, from 3650 A to 7200 A.
Physical intensities equalized.

1. Leptinotarsa decemlineata Say (Col.). 372 larvse, 1 to f grown.

Four tests.

2. Lophyrus lecontei Fitch (Hymen.). 124 larvse, full grown. Two

tests.

3. Macremphytus sp. (Hymen.). 491 larvse, full grown. Three tests.

Larvse starved.

4. Diacrisia virginica Fab. (Lep.). 92 larvse, full grown. Three tests.

5. Dryocampa rubicunda Fab. (Lep.). 100 larvse, full grown. Three

tests.

6. Anisota senatoria A & S (Lep.). 125 larvse, i to f grown. Three

tests.

Figure 4. Behavior of lepidopterous larvse to 10 wave-length bands, in dis-
array, from 3650 A to 7200 A. Physical intensities equalized.

1. Eudamus tityrus Fab. 125 larvse, full grown, starved. Three tests.

2. Hadena turbulenta Hbn. 263 larvse, f grown. Three tests.

3. Datana integerrima G. & E. 188 larvse, f grown, starved. Two tests.

4. Datana ministra Dru. 303 larvse, f to full grown. Three tests.

5. Melalopha inclusa Hbn. 70 larvse, full grown. Two tests.

6. Hyparpax aurora S. & A. 274 larvse, full grown. Three tests.
Figure 5. Behavior of lepidopterous larvse to 10 wave-length bands, in dis-
array, from 3650 A to 7200 A. Physical intensities equalized.

1. Actias luna Linn. 63 larvse, | grown. Two tests.

2. Telea polyphemus Cram. 141 larvae, f-full grown. Two tests.

3. Telea polyphemus Cram. 136 larvse, full grown, starved. Four tests.

4. Ceratomia catalpce Bdv. 110 larvse, full grown. Three tests.

5. Phlegethontius Carolina Linn. 32 larvse, full grown. Two tests.

(Plate VI)

(Jour. N. Y. Ent. Soc.), Vol. LII

7200 A

Mar., 1944]

Alexander: Crane-Flies

RECORDS AND DESCRIPTIONS OF NEOTROPICAL
CRANE-FLIES (TIPULID^, DIPTERA), XVII

By Charles P. Alexander
Amherst, Massachusetts

The previous instalment under this general title was published
in September, 1943 (Journal of the New York Entomological
Society, 51(3): 199-212). The materials here considered are
all from Ecuador where they were taken by Mr. William Clarke-
Macintyre, Mr. David B. Laddey, and Professor F. Martin Brown.
Some of the most interesting of the new species were taken at the
station “Zumbi,” in the Province of Santiago-Zamora, southern
Oriente, by Mr. Laddey. For a brief discussion of this station,
the preceding instalment should be consulted. I am greatly
indebted to the collectors for the privilege of retaining the types
of the novelties in my collection of these flies.

Genus Gnophomyia Osten Sacken

Gnophomyia (G-nophomyia) argutula new species.

General coloration of mesonotal praescutum gray with three dark brown
stripes, the lateral portions orange-yellow; thoracic pleura variegated dark
brown and obscure yellow; knobs of halteres dark brown; femora yellow, the
tips conspicuously blackened; wings yellowish gray, with three darker clouds
or bands, the second at the level of cord; cell 1st M2 widened outwardly,
nearly four times as wide at outer end as at base; cell 2nd A broad; abdom-
inal segments bicolored, the basal portions reddish yellow, the remainder
dark brown ; male hypopygium with the outer dististyle acute at tip ; phallo-
some unusually broad and obtuse.

Male. — Length about 7 mm.; wing 7.5 mm.

Rostrum brown; palpi black. Antennae brownish black; flagellar segments
nearly cylindrical; verticils of outer segments much longer and more con-
spicuous than those of the basal segments. Anterior vertex dull orange, rela-
tively narrow, the eyes correspondingly large and protuberant; posterior
portion of head brownish gray, the anterior orbits obscure orange.

Pronotum obscure brownish yellow above, dark brown on sides ; pretergites
obscure yellow. Mesonotal praescutum with the ground color of interspaces
gray, with three entire dark brown stripes, the median one more reddened at
cephalic end; humeral and lateral portions of sclerite obscure orange yellow;
posterior sclerites of notum dark brown, sparsely pruinose; scutellum paler,

Journal New York Entomological Society

[Vol. Lll

with a central, dark brown spot ; dorsal pleurotergite obscure yellow. Pleura
chiefly dark brown, sparsely pruinose, restrictedly but conspicuously varie-
gated with obscure yellow, distributed as follows: Dorsal portion of sterno-
pleurite ; posterior border of pteropleurite, and meral region. Halteres short,
stem yellow at base, the remainder dark brown. Legs with the coxae brown,
sparsely pruinose; trochanters obscure yellow; femora yellow, the tips rather
broadly and conspicuously blackened; tibiae and tarsi yellow, only the ter-
minal segment weakly darkened. Wings with the ground color yellowish
gray, the color greatly restricted by three more or less distinct, slightly
darker clouds or bands, the most conspicuous at the cord and over outer end
of cell 1st M2; slightly less distinct bands at proximal fourth of wing and
as a nearly apical darkening in the cells beyond cord; stigma very long and
narrow, dark brown; veins yellow in the ground areas, brown in the darkened
fields. Venation: Sc± ending about opposite the short transverse R2’, -Ss in
longitudinal alignment with E5, r-m at its fork; cell 1st M2 strongly widened
outwardly, nearly four times as wide at outer end as at base; cell 1st M2
approximately as long as vein M± beyond it; cell 2nd A noticeably shorter
and broader than in duplex.

Basal abdominal segments bicolored, dark brown, the basal rings conspicu-
ously reddish or reddish yellow, the subterminal segments more uniform dark
brown; hypopygium and preceding segment more yellowish; sternal pattern
generally like the tergal. Male hypopygium with the outer dististyle much
less conspicuously flattened than in laticincta, its apex acute; base of style
with only two elongate setae. Inner dististyle about one-half as long as the
outer style, provided with numerous setae, including about three of unusual
length. Phallosome unusually broad and obtuse.

Holotype, J1, Zumbi, Rio Zamora, Santiago-Zamora, altitude
700 meters, November 1, 1941 (Laddey).

The nearest relatives are Gnophomyia ( Gnophomyia ) duplex
Alexander and G. (G.) laticincta Alexander, which have the legs
and wings somewhat similarly patterned. The former species is
still known only from the female sex, differing from the present
fly in the venation and in the details of coloration of body and
wings. The latter species, laticincta , differs conspicuously in
the structure of the male hypopygium, especially of the outer
dististyle.

Gnophomyia (Gnophomyia) bulbibasis new species.

General coloration of mesonotum dark brown, very sparsely pruinose, on
praeseutum forming a discal shield; thoracic pleura striped longitudinally
with dark brown and reddish; halteres darkened; legs pale brown; wings
grayish subhyaline, stigma scarcely differentiated; male hypopygium with
the outer dististyle conspicuously bulbous just beyond base; gonapophyses.
appearing as blackened spines.

Mar., 1944]

Alexander: Crane-Flies

Male. — Length about 4.5-5.5 mm. ; wing 5-6.2 mm.

Female. — Length about 5 mm. ; wing 5.5 mm.

Rostrum and palpi dark brown. Antennae dark brown; basal flagellar seg-
ments subcylindrical, the outer ones shorter; verticils subequal in length to
the segments. Head dark gray; eyes large; anterior vertex only a little
wider than the diameter of scape.

Pronotum above yellow, darker on sides. Mesonotum almost uniformly
dark brown, very sparsely pruinose, on prsescutum forming a discal shield
that leaves the humeral and lateral portions yellowish, in cases more obscure
than in others. Pleura reddish, with a conspicuous dark brown dorsal stripe
extending from the propleura to the postnotum, passing above the halteres;
dorsopleural region yellow, confluent with the similarly colored lateral
prsescutal borders; immediately ventrad of the dark pleural stripe a more or
less distinct paler longitudinal line extending from behind the fore coxae
to the base of abdomen. Halteres dusky, the knob still darker. Legs with
coxae obscure yellow to testaceous yellow; trochanters yellow; remainder of
legs pale brown, the femoral bases clearer yellow; outer tarsal segments pass-
ing into darker brown. Wings grayish subhyaline, the extreme base yellow ;
stigmal area very restricted and pale, scarcely differentiated; veins pale
brown, yellow in the prearcular field. Venation: Sc long, S cx ending just
before level of B2, Sc2 some distance from its tip, lying opposite or before
the fork of Bs; r-m before or close to fork of Bs-} m-cu about three-fourths
its own length beyond the fork of M .

Abdominal tergites brownish black, the sternites a trifle more piceous;
hypopygium yellowish brown. Ovipositor with eerci relatively short and
stout, with setae to the tips of the valves. Male hypopygium with the outer
dististyle conspicuously expanded or bulbous just beyond base, thence nar-
rowed to a long straight rod that terminates in an acute point. Inner disti-
style much shorter, obtuse at tip; at base with about four powerful setae,
with other shorter setae on distal half, chiefly near apex. Gonapophyses
appearing as blackened spines.

Holotype, Zumbi, Rio Zamora, Santiago-Zamora, altitude
700 meters, November 4, 1941 (Laddey). Allotopotype, 5? with
the type. Paratopotypes, 2 1 $, October 31-November 2,

1941 (Laddey).

Gnophomyia ( Gnophomyia ) bulbibasis has the structure of
the male hypopygium, especially of the gonapophyses, somewhat
as in G. (G.) oxymera Alexander, from which it differs in other
hypopygial characters, as the bulbous basal enlargement of the
outer dististyle.

Gnophomyia (Gnophomyia) fessa new species.

General coloration of mesonotum and the dorsal pleurites dark brown,
ventral pleurites reddish brown ; rostrum, antennee and halteres black ; femora

Journal New York Entomological Society

[Vol. LII

obscure yellow, with a narrow subterminal darker ring; wings subhyaline,
stigma reduced to a narrow seam; male hypopygium with the inner dististyle
bulbous, the apex conspicuously wider than the base; gonapophyses incurved,
appearing as separate blades, blackened and microscopically serrulate at
bases.

Male. — Length about 5 mm.; wing 5 mm.; antenna about 1.5 mm.

Female. — Length about 6.5 mm. ; wing 6 mm.

Rostrum and palpi brownish black. Antennae black throughout; basal
flagellar segments subcylindrical, the outer ones more elongate-oval, with
verticils that exceed the segments in length. Head dark gray; eyes (male)
relatively large, the anterior vertex correspondingly narrowed.

Pronotum brown, the pretergites conspicuously light yellow. Mesonotum
almost uniform dark brown, the surface very sparsely pruinose, the central
portion of praescutum and the scutal lobes slightly darker. Pleurotergite and
dorsal pleura dark brown, contrasting abruptly with the reddish brown
ventral pleurites, the surface sparsely pruinose. Halteres brownish black
throughout. Legs with coxae reddish; trochanters yellow; femora obscure
yellow, more infuscated immediately before their tips to form a narrow,
nearly terminal ring; tibiae and basitarsi obscure yellow, the tips narrowly
infuscated; outer tarsal segments passing into brownish black. Wings sub-
hyaline, with a very faint darker tinge; stigma darker brown but reduced
to a linear area adjoining veins B^ and B1+2, scarcely involving the surround-
ing membrane; prearcular and costal fields a trifle more yellowish; veins
brown, somewhat more brightened in the basal areas. Venation : $<?! ending
opposite -K2; Bs straight, oblique; basal section of B5 lacking, r-m at fork
of Bs; B2 variable in position, before the fork of B2+ 3+4 or nearly its own
length beyond this fork; cell 1st M2 narrow, subequal in length to vein
beyond it; m-cu from one-third to two-thirds its length beyond the fork of M.

Abdominal tergites and hypopygium brownish black; sternites yellow.
Ovipositor with cerci relatively small, only weakly sclerotized, with setae
virtually to their tips. Male hypopygium with the outer dististyle a simple,
curved, relatively narrow, blackened rod, the tip subacute. Inner dististyle
relatively short, bulbous, the apex almost twice as wide as the base, with
several setae, chiefly on the distal third. Phallosome with the gonapophyses
incurved, appearing as separate blades, at apex blackened and microscopically
roughened to serrulate; apex of phallosome narrow.

Holotype, Palmar, Rio Maizito, Manabi, altitude 200 meters,
May 20, 1941 (Laddey). Allotopotype, §, pinned with type.

Gnophomyia ( Gnophomyia ) fessa is most similar to species
such as G. (G.) oxymera Alexander and G. ( G .) nectarea new
species, differing in the details of coloration and, especially, the
structure of the male hypopygium, as the inner dististyle and
the phallosome.

Mar., 1944]

Alexander: Crane-Flies

G-nophomyia (G-nophomyia) nectarea new species.

Size small (wing, male, about 5 mm.) ; mesonotum and dorsal pleura
opaque black, the ventral pleurites abruptly reddish; halteres dusky; legs
brownish yellow; wings subhyaline, the stigma and an extensive cloud on
proximal third of wing infuscated; vein R2 very faint to nearly atrophied;
cell 1st M2 long and narrow ; cell 2nd A relatively narrow ; male hypopygium
with the outer dististyle. slender; inner dististyle with a conspicuous basal
lobe that is microscopically corrugated or wrinkled.

Male. — Length about 4-4.8 mm.; wing 4.5-5.5 mm.; antenna about 1.3-
1.6 mm.

Female. — Length about 5 mm. ; wing 5 mm.

Eostrum dark brown; palpi black. Antennae of moderate length, dark
brown; flagellar segments subcylindrical ; longest verticils subequal in length
to the segments. Head brown, the orbits and posterior vertex light gray;
anterior vertex relatively narrow, about one-third the diameter of scape; eyes
(male) correspondingly large.

Pronotum dark brown ; pretergites light yellow. Mesonotum chiefly
blackened, the surface opaque by a sparse pruinosity; central portion of
scutum and posterior border of scutellum slightly more reddened. Dorsal
pleurites covered by a broad black longitudinal stripe, this area also involv-
ing the pleurotergite and surrounding the root of halteres; ventral pleurites
reddish, very sparsely pruinose. Halteres dusky, base of stem restrictedly
brightened. Legs with coxae and trochanters yellow; remainder of legs
brownish yellow, the outer tarsal segments more infuscated. Wings sub-
hyaline, the extreme base restrictedly yellow; stigma oval, darker brown;
a conspicuous dusky cloud or wash on basal third of wing in general vicinity
of vein Cu, involving the bases of cells Cu, 1st A and 2nd A, together with
much of M ; veins brown, yellow in the flavous basal region. Venation:
Vein R2 very faint to nearly atrophied; Rs in direct longitudinal alignment
with _K5; branches of Rs all extending generally parallel to one another;
cell 1st M2 relatively long and narrow, its inner end pointed or strongly nar-
rowed, the outer end more widened, the cell subequal in length to vein M4;
m-cu less than its own length beyond fork of M, at near one-fourth to one-
fifth the length of the cell; cell 2nd A relatively narrow.

Abdominal tergites and the hypopygium brownish black; basal sternites
a trifle paler. Male hypopygium with the outer dististyle unusually slender
throughout, narrowed at apex to a subacute point. Inner style with a con-
spicuous basal lobe or shoulder, its surface and adjoining margin of style
microscopically wrinkled. Phallosome of moderate width, its tip blackened
and slightly narrowed to an obtuse point, the surface microscopically wrinkled.

Holotype, J1, Zumbi, Rio Zamora, Santiago-Zamora, altitude
700 meters, November 2, 1941 (Laddey). Allotopotype, §•
Paratopotypes, 2 jy1, October 30-November 5, 1941 (Laddey).

Journal New York Entomological Society

[Vol. LII

Gnophomyia ( Gnophomyia ) nectar ea is entirely distinct from
other small-sized regional members of the genus. From all such,
it differs conspicuously in the peculiar wing pattern and in the
structure of the male hypopygium.

Gnophomyia (Gnophomyia) tuber new species.

General coloration of mesonotum medium brown, with poorly defined pat-
tern; pleura with a broad, medium brown, longitudinal stripe over the dorsal
sclerites; halteres yellow; wings with a buffy tinge, brighter in the basal
portions; abdominal tergites reddish brown, weakly darkened laterally;
male hypopygium with the basistyles short, on mesal face near base with a
conspicuous darkened tubercle, this provided with about three conspicuous
setae; outer dististyle compressed-flattened, its apex obtuse; inner dististyle
with the apex very obtuse to subtruncate.

Male. — Length about 6-6.3 mm.; wing 6.8-7 mm.; antenna about 1.5-
1.6 mm.

Rostrum testaceous brown; palpi brownish black. Antennae with the scape
and pedicel brownish yellow, flagellum brownish black; flagellar segments
long-cylindrical, the terminal segments shorter; verticils considerably exceed-
ing the segments in length. Head brownish gray; anterior vertex relatively
narrow, eyes large.

Pronotum light yellow above, darker on sides; pretergites light yellow.
Mesonotal praescutum medium brown, sparsely pruinose, the median area
more darkened, behind forming more or less distinct stripes; scutal lobes
similarly darkened, the posterior border obscure yellow; scutellum brown,
with a yellow spot on either side of base; postnotum medium brown, more
yellowish on suture between mediotergite and pleurotergite. Pleura with a
broad but relatively inconspicuous, medium brown stripe, more intense in
front, becoming diffuse behind; dorsopleural region yellow; ventral pleurites
obscure yellow, paler yellow behind. Halteres yellow. Legs with the coxae
and trochanters yellow; femora obscure yellow, weakly darkened at or near
tips; tibiae and basitarsi yellow, the tips narrowly and inconspicuously infus-
cated; outer tarsal segments brownish black. Wings relatively broad, with
a sandy or buffy tinge, the prearcular and costal fields clearer yellow ; stigma
very restricted in area, pale brown; veins brown, more brownish yellow in
the brightened portions. Venation: Scx ending about opposite the fork of
E2+3+i, £u2 nearly opposite the fork of Es ; E2+3 more than one-third E2+3+i;
Es in direct longitudinal alignment with E5 ; r-m beyond fork of Es ; cell
1st M2 relatively long, subequal to vein Jf4 beyond it ; m-cu about two-thirds
its length beyond fork of M.

Abdominal tergites reddish brown, weakly darkened laterally; sternites
clearer yellow; hypopygium yellowish brown. Male hypopygium with the
basistyles short, on mesal face near base with a conspicuous darkened tuber-
cle, this provided with about three conspicuous setae. Outer dististyle dark-
ened, conspicuously compressed-flattened, widest at near midlength, its apex

Mar., 1944]

Alexander: Crane-Flies

obtuse. Inner dististyle extending about to midlength of the outer style,
dark-colored, its apex very obtuse to subtruncate; rather numerous setae,
including a row of four or five larger ones along the lower or cephalic margin.
Phallosome relatively broad, the apex obtuse.

Holotype, J', Playas de Montalvo, Los Rios, altitude 15 meters,
March 5, 1938 (Macintyre). Paratopotypes, 1 <$, 1 sex?, pinned
with type.

In its general appearance and wing coloration, the present fly
is most like Gnophomyia ( Gnophomyia ) acricula Alexander and
G. ( G .) digitiformis Alexander, but has a very different hypo-
pyginm. The tubercle on the basistyle is not found in any other
species known to me.

Genus N eo gnophomyia Alexander

Neognophomyia spectralis new species.

General coloration of mesonotal praescutum reddish yellow, the posterior
sclerites darker; vertex yellow, with a brown central area; thoracic pleura
with a conspicuous dorsal black stripe; tips of femora, tibiae and basitarsi
broadly blackened; wings with a pale yellow tinge, patterned with brown,
including a subbasal fascia; abdominal tergites conspicuously patterned
with brownish black and yellow; male hypopygium with the tergal spines
blackened, nearly straight; phallosome produced at apex into lateral horns.

Male. — Length about 5-5.5 mm.; wing 6-7 mm.

Rostrum obscure yellow; palpi brown. Antennae with scape and pedicel
dark brown; basal flagellar segments pale brown, the outer ones paling to
brownish yellow; flagellar segments passing from oval through long-oval;
verticils conspicuous. Head yellow, the anterior vertex with a conspicuous
brown area; anterior vertex moderately wide, a little more than twice the
diameter of the scape.

Pronotum yellow medially, brownish black on sides. Mesonotal praescutum
reddish yellow, more yellowish on lateral and humeral portions, in cases more
darkened, especially near suture; scutum yellow, each lobe with two brown
areas; scutellum black, more or less pruinose; postnotum brownish black,
including both the mediotergite and pleurotergite. Pleura reddish yellow
with a broad black longitudinal stripe beginning on sides of pronotum, in-
volving the anepisl;ernum, dorsal pteropleurite and the postnotum, as de-
scribed. Halteres yellow. Legs with the coxae yellow to reddish yellow;
trochanters yellow; femora, tibiae and basitarsi yellow, with broad and
conspicuous black tips; remainder of tarsi black. Wings with a pale yellow
tinge, patterned with brown, including a broad seam from stigma across
anterior cord, and narrower seams on m-cu and outer end of cell 1st M2; a
less conspicuous subbasal band from origin of Rs extending obliquely across
cell M into cells Cu, 1st A and 2nd A ; extreme base of wing blackened; veins

Journal New York Entomological Society

[Vol. LII

yellow to brownish yellow, darker brown in the patterned areas. Venation:
B2 at near midlength of petiole of cell _E3; cell 1st M2 strongly narrowed at
proximal end; m-cu from about one-half to approximately its own length
beyond the fork of M.

Abdominal tergites yellow, handsomely patterned with brownish black,
the segments chiefly darkened with a large yellow area at posterior border
of each segment, encroaching on the base of the succeeding segment, the
outer segments more uniformly darkened, greatly restricting the yellow color ;
sternites yellow, the terminal segments more darkened; hypopygium chiefly
obscure brownish yellow. Male hypopygium with the tergal spines appearing
as long, nearly straight, blackened blades, a little expanded at base, thence
gradually narrowed to the acute tips. Outer dististyle relatively slender;
outer margin of distal third with five long setae, additional to the single
terminal bristle. Inner dististyle large and massive, terminating in a broad
blackened beak, the apical border with about seven or eight strong setae;
basal tooth or flange of style strongly blackened, provided with four strong
setae. Phallosome relatively wide, at apex produced into a slender spinous
point that is directed laterad, immediately cephalad of which is a broadly
rounded emargination.

Holotype, J1, Banos, Tungurahua, altitude 2,000 meters, July
14, 1939 (Macintyre). Paratopotypes, 7 altitude 1,700-
2,000 meters, May 11-June 2, 1937 ; paratype, 2 J'J1, Pititi, near
Banos, altitude 1,900 meters, June 14, 1937 (Macintyre).

The only generally similar species is Neognophomyict hirsuta
(Alexander) of eastern Brazil, which has the pattern of the
body, legs and wings somewhat the same but with the male
hypopygium entirely different. This latter species has the sub-
basal darkened wing band much wider and more continuous than
in the present fly.

Neognophomyia interrupta new species.

Allied to hostica; general coloration yellow, the praescutum with the disk
chiefly reddish yellow ; wings grayish yellow, with a narrow brown seam along
cord; abdomen pale, with tergites three, five and six conspicuously dark
brown; male hypopygium with the outer dististyle strongly constricted at
near midlength, the bulbous outer portion terminating in two subequal elon-
gate setae; tergal spines pale, angularly bent beyond midlength.

Male. — Length about 4.5 mm.; wing 5 mm.

Eostrum pale yellow; palpi pale, the outer segments darkened. Antennae
with scape and pedicel light yellow; flagellum broken. Head obscure brown-
ish yellow; eyes (male) relatively large.

Pronotum yellow. Mesonotal praescutum yellow, with three more or less
confluent more reddish stripes that form a nearly continuous discal area;

Mar., 1944]

Alexander: Crane-Flies

scutal lobes brownish black, the remainder of scutum more brownish yellow ;
scutellum and mediotergite brownish yellow, the pleurotergite almost covered
by an oval velvety black spot, as is common in the genus. Pleura reddish yel-
low the anepisternum a little infuscated. Halteres uniformly pale yellow.
Legs with the coxae and trochanters yellow ; remainder of legs broken. Wings
grayish yellow, the prearcular and costal fields somewhat brighter yellow;
a narrow but conspicuous brown seam along cord, becoming indistinct or
obliterated at fork of M ; veins brownish yellow, darker in the infuscated
areas. Venation: E2 at about one-third the length of petiole of cell R3;
vein ^3‘ oblique; cell 1st M2 only slightly widened outwardly; m-cu at near
one-third the length of the cell.

Abdomen conspicuously patterned; basal tergites brownish yellow, dark-
ened laterally; tergites three, five and six conspicuously dark brown; tergites
four, seven and eight pale; hypopygium and sternites more uniformly yellow.
Male hypopygium with the outer dististyle strongly constricted at near mid-
length, the base widened, the apex bulbous, its extreme tip broadly obtuse
with two subequal elongate setae. In hostica, the style is unusually slender,
gradually narrowed beyond the basal enlargement, the tip unusually slender
and thus with only a single apical seta. Inner dististyle much as in hostica,
more widened at near midlength, the setae at this point slightly more
numerous, longer and paler. Phallosome narrower. Tergal spines somewhat
as in hostica, the basal section shorter.

Holotype, J', Zumbi, Rio Zamora, Santiago-Zamora, altitude
700 meters, November 5, 1941 (Laddey).

The most similar described species is Neognophomyia hostica
Alexander, of Peru, which differs most evidently in the colora-
tion of the abdomen and in the structure of the male hypopygium,
as compared above.

Genus Gonomyia Meigen

Cronomyia (Progonomyia) acrissima new species.

General coloration gray, the praescutum with three conspicuous brown
stripes; thoracic pleura dark gray with a broad yellow longitudinal stripe;
legs dark brown, the tarsi black; wings with a weak grayish tinge, unpat-
terned except for the very pale brown and inconspicuous stigma; male
hypopygium with the mesal face of basistyle before apex with a longitudinal
group of acute spines, these exceeding 60 in number.

Male. — Length 5.5-5. 6 mm.; wing 6-6.2 mm.; antenna about 1-1.1 mm.

Rostrum black, sparsely pruinose; palpi brownish black. Antennae with
the scape dark brown, pedicel and flagellum black; flagellar segments sub-
cylindrical to long-oval with verticels that are subequal in length to the
segments. Head gray; vertex with a median brown line.

Pronotum blackened, pruinose, the lateral portions, together with the
anterior pretergites, obscure yellow. Mesonotal praescutum gray, with three

Journal New York Entomological Society

[Vol. LII

conspicuous brown stripes, the median one not reaching the suture, the lat-
erals crossing the suture onto the scutal lobes; pseudosutural foveae black,
conspicuous; posterior sclerites of notum gray, the posterior border of scu-
tellum more obscure yellow; postnotum gray, the dorsal portion of pleuro-
tergite and adjoining portion of mediotergite obscure yellow. Pleura dark
gray, with a broad and conspicuous yellow stripe extending from behind the
fore coxae across the dorsal sternopleurite, behind including the ventral
pteropleurite, meron and metapleura. Halteres with stem yellowish brown,
knob darker brown. Legs with coxae light brown, sparsely pruinose; tro-
chanters brownish yellow; remainder of legs dark brown, the tarsi passing
into black. Wings with a weak grayish tinge, unpatterned except for the
very pale brown stigma; extreme wing base paler; veins brown, brightened
at base of wing. Venation: Sc relatively short, Sol ending a distance be-
yond origin of Bs about equal to m-cu, Sc2 a slightly shorter distance before
origin; Bo indicated by a very faint element at midlength of stigma; cell
2nd M2 nearly twice its petiole; m-cu from three-fifths to two-thirds its
length before the fork of M.

Abdominal tergites dark grayish brown, sternites slightly paler; eighth
and ninth segments somewhat paler brown, the styli darker. Male hypo-
pygium with the apex of basistyle obtuse, unarmed; mesal face immediately
back from tip with an extensive longitudinal area of acute spines, these
totalling in excess of 60. Outer dististyle a semicylindrical blade, its apex
narrowed into an acute black spine. Intermediate style a long darkened
blade, about one-half longer than the outer style; at near one-third its length
bearing a more slender branch or arm that is approximately one-half as long
as the main apical branch; stem near the branch with numerous setae, these
becoming even longer and more conspicuous on base of lateral branch; main
or axial branch glabrous, gradually narrowed to the subacute cultriform
apex. Inner dististyle broad, obtuse at apex, near margin provided with
several setae, those at and near apex longest. JEdeagus relatively slender,
narrowed outwardly, near apex with small and inconspicuous lateral shoulders.

Holotype, Banos, Tungurahua, altitude 1,450 meters, April
23, 1939 (Macintyre). Paratopotype, 1 J*.

The present fly is readily distinguished from all other gen-
erally similar forms by the structure of the male hypopygium,
especially the spinous area near apex of basistyle. Such arma-
ture is uncommon in the subgenus and in all other cases the num-
ber, arrangement and shape of the spines is different; in
Gonomyia ( Progonomyia ) compact a Alexander there is a rela-
tively large group of spiculose points; in G. (P.) serena Alex-
ander, the spines of the general type of the present fly but very
few in number; in G. (P.) thiosema Alexander, an even larger
group of elongate spinous pegs on mesal face of basistyle but

Mar., 1944]

Alexander: Crane-Flies

these with obtusely rounded tips. In all these species, the
structure of the dististyles is likewise distinctive.

G-onomyia (Progonomyia) hyperplatys new species.

General coloration of notum grayish brown; antennae black throughout;
pleura blackened, sparsely pruinose, with a conspicuous yellow longitudinal
stripe; knobs of halteres darkened; wings yellowish, restrictedly patterned
with pale brown; S c± ending shortly beyond origin of Es-, male hypopygium
with the phallosome unusually wide, appearing as a broadly flattened plate,
each outer lateral angle further produced.

Male. — Length about 6 mm.; wing 6.6 mm.

Eostrum and palpi dark brown. Antennae brownish black throughout;
flagellar segments subcylindrical, passing into long-oval, with verticils that
exceed the segments. Head brownish gray on anterior vertex, deepening in
color behind the antennal bases, the broad posterior vertex clear gray.

Pronotum obscure yellow medially, dark brown sublaterally ; pretergites
yellow. Mesonotum dark grayish brown, the humeral and lateral portions
light yellow; scutellum more reddish brown; mediotergite dark brown, with
a conspicuous yellow area on either side of basal half; dorsal portion of
pleurotergite similarly yellow. Pleura blackened, sparsely pruinose, with
a conspicuous yellow longitudinal stripe extending from above the fore coxae
across the dorsal sternopleurite to the ventral pteropleurite and meral region.
Halteres with stem yellow, knob weakly darkened. Legs with the coxae
basally dark brownish gray, the tips restrictedly yellow; trochanters brown;
remainder of legs broken. Wings with a yellowish tinge, restrictedly pat-
terned with rather pale brown, the areas chiefly restricted to the vicinity
of the veins, distributed as follows: Origin of Es ; stigma; cord and base
of cell 2nd M2 ; two small isolated spots in Anal cells, one near outer end
of cell 1st A adjoining vein 2nd A, the other near center of cell 2nd A ;
veins brown. Venation: Sc short, S cx ending shortly beyond origin of Es,
Sc2 about an equal distance before this origin, Es relatively long, square and
spurred at origin; E2 about twice E3+i; m-cu from about one-third to one-
half its length before fork of M.

Abdomen brownish black; ninth segment abruptly yellow, the styli again
blackened. Male hypopygium with the outer dististyle a short, slender,
curved rod that narrows to the acute blackened tip, the outer margin with
delicate microscopic setulae. Intermediate style broadly flattened, entirely
dark-colored, the inner margin at near midlength produced into a sharp spine,
the surface basad of this with conspicuous setae; apical portion of style ap-
pearing as a broad flattened basal flange that narrows into a long arm that
is narrowly but deeply notched at apex. Inner dististyle a dusky lobe with
strong spinous setae along the entire inner face, the terminal one longest.
Phallosome unusually wide for a member of this subgenus, appearing as a
broadly flattened plate subtending the aedeagus, each outer lateral angle pro-
duced further into a flattened lobe or blade, with about seven setae chiefly
distributed along the outer margin near apex.

Journal New York Entomological Society

[Vol. LII

Holotype, Banos, Tungurahua, altitude 1,450 meters, April
23, 1939 (Macintyre).

The present fly is entirely different from other described species
of the subgenus, differing especially in the structure of the male
hypopygium and particularly of the phallosome. The most
similar form is Gonomyia ( Progonomyia ) velutina Alexander,
which differs in the wing pattern and in all details of structure
of the male hypopygium.

Genus Cryptolabis Osten Sacken

Cryptolabis (Cryptolabis) alticola new species.

General coloration black, more or less prninose ; lateral pretergites abruptly
yellowish white; legs black; wings with a strong brownish tinge, the pre-
arcular and costal fields yellow; Bs very long; male hypopygium with the
dististyle large and conspicuous, exserted, consisting of an outer flattened
setuliferous blade and a bispinous inner body; tergal plate bearing conspicu-
ous lateral arms that terminate in several strong setae; aedeagus long and
slender.

Male. — Length about 4.5 mm.; wing 5.2 mm.

Rostrum and palpi black. Antennae brownish black; flagellar segments
oval, with conspicuous verticils. Head black, presumably pruinose in fresh
specimens.

Pronotum, mesonotum and pleura of the unique type dull black, the surface
presumably more or less pruinose in fresh specimens; pretergites abruptly
and conspicuously yellowish white. Halteres brown, the apex of knob
brighter. Legs black, with relatively conspicuous suberect setae. Wings
with a strong brown tinge, the prearcular and costal fields yellow; axillary
region restrictedly infuscated; veins and macrotrichia brown. Macrotrichia
of cells relatively abundant, beyond the cord extending from outer end of
cell B2 to cell Mi} in the radial field involving all of the cells with the excep-
tion of the basal fourth to fifth. Venation: Sc relatively short, Sc1 ending
some distance before the end of Bs; Bs very long, exceeding in length vein
Bs; B2+ 3+4 at origin nearly perpendicular to Bs , subequal in length to B2+ 3;
cell M3 deep; m-cu at near midlength of Ms+i; vein 2nd A gently sinuous.

Abdomen brownish black; hypopygium black. Male hypopygium with the
dististyle large and conspicuous, exserted; consisting of an outer fleshy lobe
or blade that may represent a separate style, and the main body of the style
itself; outer blade flattened, densely covered with short setae; inner portion
of style with apex dilated into a truncated portion, the outer margin with
two strong blackened spinous points, the more basal one erect, the outer spine
more appressed. What appears to be the tergite appears as a transverse
plate with the median portion strongly produced into a rounded lobe, the
lateral arms appearing as strong cylindrical lobes that are directed strongly

Mar., 1944]

Alexander: Crane-Flies

mesad and then caudad, the tips with about seven elongate setae. .ZEdeagus
unusually long and slender, blackened, transversely corrugated.

Holotype, J1, Hacienda Talahua, Bolivar, altitude 3,100 meters,
April 28, 1939 (Brown).

Cryptolabis ( Cryptolabis ) alticola is entirely different from the
other described species of the genus. It is most similar to species
such as C. (C.) chilotanica Alexander, of southern Chile, yet very
distinct in the structure of the male hypopygium. For a discus-
sion of the type locality, consult Brown (Ann. Ent. Soc. Amer.,
34: 848; 1941).

Mar., 1944]

Clench: Lyc^enhle

TWO NEW SUBSPECIES OF EVERES
COMYNTAS GODART (LEPIDOPTERA,
EYC/ENIDJE)

By Harry K. Clench
Cambridge, Mass.

Two races of this wide-ranging species have recently come to
my attention, one from South Dakota and one from Montana.

Everes comyntas valerise, new subspecies

TJpperside :

Male. Both wings slightly purplish blue. Fore wing with a narrow dark
border on the outer margin. Costa and costal veins pencilled with light blue.
Hind wing with an internervural row of small spots on the outer margin.
Costa dark bordered. Cu2 with a short tail. Fringe of fore wing dark
basally, white outwardly; of the hind wing white.

Female. Both wings brown. Base of each dark blue. Hind wing with a
black spot in the CurCu2 interspace basally bordered by a shallow, rather
dull orange lunule. A thin marginal pale bluish line borders the outer mar-
gin, interrupted at the veins, basal to which is a similar but scalloped and
more obscure line. The two outline a series of internervural dark spots.
Occasionally these lines are very faint, and the row of spots consequently
almost indistinguishable.

Underside :

Male. Ground color grayish tan. All spots arranged as in typical comyn-
tas, but with the post-discal series usually rounder and darker — occasionally
very heavy. The orange over the two spots in the M3-Cux-Cu2 interspaces of
the hind wing is usually faint, although stronger in the latter than in the
former. On this same wing, just basal to the marginal compound border,
the ground color is white between the veins.

Female. Similar to the male.

Length of fore wing: Male, 11.5-13.5 mm.; Female, 10-12.5 mm.

Holotype, male, near Lead, South Dakota, June 22, 1939 v(V.
H. and A. C. Frederick).

Allotype, female, Terry Peak, South Dakota, el. 5200+ feet,
June 24, 1939 (V. H. and A. C. Frederick).

Paratypes, 1 male, same data as holotype ; 1 male and 2 females,
same data as allotype; 1 female, Harney Peak, el. 5200+ feet,
June 25, 1939 (A. C. Frederick) ; 31 males, 3 females, Spearfish
Canyon and vicinity, el. 5200+ feet, as follows : 3 males, June 26,

Journal New York Entomological Society

[Vol. Li l

27, and July 1, 1939, resp. (V. H. and A. C. Frederick) ; 15 males,
2 females, June 29, 1942 (A. C. Frederick) ; 13 males and 1
female, June 30, 1942 (A. C. Frederick) ; 17 males, Icebox Can-
yon, June 29, 1942 (A. C. Frederick). All localities in the Black
Hills, South Dakota.

Holotype and allotype no. 25934 in the collection of the Museum
of Comparative Zoology. Paratypes in the collection of Mr.
Frederick and of the author.

Remarks. This subspecies appears to be most similar to the
southwestern race herrii Grinnell1 from which it differs in the
following particulars : the black border on the wings above in the
male is slightly thinner. This is most apparent on the hind wing,
where the border in herrii is thick enough to include the inter-
nervural spots, while in valerice these spots are almost always
free. There is no orange lunule on the hind wing above, an
almost constant feature of herrii , and even more prominent in
typical comyntas, but more variable in the latter. The ground
color below in both sexes appears to be a little darker. The
female of valerice is largely blackish brown above, with a rather
dark basal blue shading, while in herrii both wings are rather
extensively blue above. The orange lunules above are smaller in
females of valerice than in those of herrii.

This subspecies appears to be quite variable. One male has a
tiny orange lunule in the Cui-Cu2 interspace above — the only
indication of it in the whole type series. The same specimen and
one or two others have the marginal border on both wings thick-
ened, that on the hind wing including the submarginal spots.
Below, the intensity of the spots varies considerably. One speci-
men has the submarginal series of lunules in the compound border
enlarged and very dark, giving the insect a most peculiar appear-
ance. The post-discal series of spots may be enlarged or reduced.

This subspecies is named for Mrs. A. C. Frederick, who mate-
rially assisted her husband in collecting the type series.

Everes comyntas albrighti, new subspecies

TJpperside :

Male. Uniform violet-blue on both wings. Fore wing with a very narrow
dark marginal border. Hind wing also with this border, and in addition,.

i Can. Ent., 33 : 192, 1901.

Mar., 1944]

Clench : Lyc^enidhs

an almost obsolete row of small dashes, the heaviest in the CivCuo inter-
space.

Female. Uniform brown on both wings. Base of fore wing blue. On
the hind wing, in the Cux-Cm, interspace, a faint orange lunule surmounts a
tiny dark spot.

Underside:

Male. Fore wing dirty white, with a broad costal border of dark grayish
scaling. The marginal compound border is almost obsolete, save for a few
dark scales. A post-discal row of black spots, large and distinct, runs from
costa to inner margin, the costal ones usually obsolete. Cell closed by a dark
dash. Hind wing with a brownish gray ground color, fading on the outer
margin. The submarginal border is, as in the fore wing, almost completely
absent. The post-discal series, the cell-end bar, and the basal spots are also
nearly gone, but their positions are shown by whitened areas in the gray.
In the Cuj-Cuo interspace is a small black spot, obscurely scaled with metallic,
and capped by a tiny orange crescent.

Female. The single female examined shows a very similar appearance.
The post-discal series of spots on the fore wing is represented, however, only
by spots from M;> to the inner margin. The costal gray-brown on this wing
extends down to M3.

Length of fore wing: Male, 11-11.5 nun.; female, 11 mm.

Holotype, male, Kings Hill, Montana, July 9, 1939 (from C. C.
Albright).

Allotype, female, same data.

Paratypes, two males, same data.

Holotype and allotype to be deposited in the collection of the
Museum of Comparative Zoology. One paratype in the collec-
tion of Mr. C. F. dos Passos. One paratype in the collection of
the author.

Remarks. This subspecies differs from all other North Ameri-
can Everes thus far known in the grayish costal shading on the
fore wing below, and in the gray ground color of the hind wing
below. This subspecies appears to be closer in appearance to
northern specimens of amyntula Boisd.2 since several of its char-
acters correspond quite closely to that species, such as the narrow
margin above, reduced maculation below (aside from the gray
ground color) and reduced orange in both sexes.

This subspecies is named for Dr. C. C. Albright, of Great
Falls, Montana, from whom the specimens were obtained.

2 Ann. Soc. Ent. France, 10 (2) : 294, 1852.

(Jour. N. Y.

Ent. Soc.), Vol. LIT

(Plate VII)

FRANK EUGENE LUTZ

Mar., 1944]

Weiss: Frank E. Lutz

FRANK EUGENE LUTZ
1879-1943

Frank E. Lutz, the son of Martin P. Lutz and Anna Amelia
(Brockway) Lutz, was born in Bloomsburg, Pa., on September 15,
1879. His early education took place in the public schools and
the Bloomsburg State Normal School. From Haverford College
in 1900 he received his A.B. degree. During his first two years
in college he specialized in mathematics, upon the advice of his
father, who, being an insurance agent, was impressed by the
large earnings of life insurance actuaries. However, after two
years the boy decided to go into medicine and so mathematics
was dropped for biology. With college over his biology teacher,
H. S. Pratt, advised him to go into biometry in view of his train-
ing in both mathematics and biology. This he did by going to see
Dr. C. B. Davenport of the University of Chicago, who had charge
of a summer biological laboratory at Cold Spring Harbor. At
this laboratory Frank E. Lutz waited on tables and counted the
grooves on scallop shells, finally publishing his first paper, a very
short one, entitled “A Study in the Variations in the Number of
Grooves upon the Shells of Pecten irradiens (Lam.) ” in Science
in 1900. Although brief, this paper helped him to get a scholar-
ship at the University of Chicago where he obtained his A.M. in
1902. While working, as biologist for the North Shore Improve-
ment Association, in mosquito control on the north shore of Long
Island, he earned enough money to go to London (Eng.) where he
studied under Karl Pearson. From September 1902 to June 1903
he was a student in London and Berlin.

From 1904 to 1909 he was employed as resident investigator at
the Station for Experimental Evolution (Carnegie Institution) at
Cold Spring Harbor, N. Y., where he did research work on hered-
ity. Within this period, or in 1907, he obtained his Ph.D. from
the University of Chicago for his dissertation on “The Variation
and Correlation of Certain Taxonomic Characters of Gryllus,”
and at the beginning of this period, or on December 30, 1904, he
married Martha Ellen Brobson, of Philadelphia, Pa.

Journal New York Entomological Society

[Vol. Lll

Dr. Lutz then entered the employ of The American Museum
of Natural History in 1909 as assistant curator in the department
of invertebrate zoology. From 1917 to 1921 he served as asso-
ciate curator and in 1921, when the department of entomology was
created, he was appointed curator. For 22 years, or until his
death at the age of 64 on the morning of November 27, 1943, at
Harkness Pavilion, New York City, after an illness of several
weeks, Dr. Lutz continued as chairman and curator of the depart-
ment of insects and spiders.

After coming to the American Museum of Natural History his
activity in biometrics declined and was replaced by an absorbing
interest in insects, although he never had any college training in
entomology and although several of his early museum papers
dealt with the history of Antarctic explorations and with the
string-figures of Patamana Indians. Under Dr. Lutz’s leader-
ship a large exhibition and study collection of insects was assem-
bled, now numbering approximately 2,000,000 specimens. Many
of these collections were made by Dr. Lutz during the course
of 23 field expeditions to various parts of the United States and
to South and Central America and the West Indies. These
expeditions started in 1908 with a trip to Cuba and Mexico, and
after he entered the employ of the Museum, 23 expeditions were
made. The first took place in 1911 to the West Indies, British
Guiana and Florida, and the last in 1941 to California. Between
these dates, Dr. Lutz collected and made observations in Florida,
Louisiana, Texas, California, Colorado, Wyoming, Utah, Idaho,
Panama, Porto Rico, Cuba and British Guiana. In all five trips
were made to the West Indies, five to Panama, five to Florida and
eight to the western part of the United States. Some of these
western trips were made in a special Museum truck-like auto-
mobile which Dr. Lutz had outfitted with equipment for collecting
and living out-of-doors, making him independent of hotels and
trains. In addition, he took an active part in Museum affairs and
commttees and was chairman of publications and editor of the
Museum’s Bulletin and Memoirs from 1917 to 1929. From 1925
to 1928 he directed the Station for the Study of Insects at Tuxedo,
N. Y., and was one of the nation’s leading exponents for nature
trails and museums.

Mar., 1944]

Weiss: Frank E. Lutz

During the summer of 1926 he began the first trailside museum
of its kind at Bear Mountain, N. Y., and was called upon by many
organizations for advice in establishing similar trails in parks and
wild-life areas in various parts of the United States. I distinctly
recall his enthusiasm upon this subject, when in company with
Mr. E. L. Dickerson I paid him a visit at Tuxedo when the sta-
tion there was half completed. Dr. Lutz was so anixous to get
things finished that he would not go with us for lunch, preferring
to dine quickly upon some pieces of bread over which he had
broken a raw egg, a nutritious, if not appetizing mixture.

In planning the arrangement of the insect exhibits at the
Museum, Dr. Lutz attempted not only to supply information
about insects, but to interest the visitor in entomology as well.
About 1915 the plan of the Hall of Insect Life involved exhibits,
with continuity, covering ontogeny, anatomy, physiology, tax-
onomy, phylogeny, life-histories, insect associations, insect
enemies and evolution. And in addition, there were exhibits on
miscellaneous topics such as insect architecture, insects as food,
medicine, social insects, etc., etc. Later as exhibit methods
changed there were originated by Dr. Lutz various habitat groups.
Dr. Lutz tells of the early days of his department in ‘‘Natural
History,’ ’ May-June, 1924, under the title “Amateur Entomolo-
gists and the Museum.” During his administration the insect
collection was increased by gifts, purchases and expeditions, well
over a million specimens.

For many years Dr. Lutz took an active part in the affairs of
the New York Entomological Society, serving as president in 1925
and 1926 and on the Publication Committee for 20 years. For a
long period the meetings were held in Dr. Lutz ’s room on the third
floor of the Museum and there, surrounded by preserved spider-
webs, Dr. Lutz’s zoo of living insects and entomological books and
paraphernalia, many interesting entomological discussions took
place, in which he always participated.

A glance at Dr. Lutz’s published writings indicates that from
1910 on, they were concerned exclusively with insects, and spiders,
principally the former. In both his popular and scientific writ-
ings he covered such topics as geographic distribution, insect
sounds, a study of ultraviolet in relation to flower- visiting habits

Journal New York Entomological Society

[Vol. LII

of insects, wind and the direction of insect flight, insect life in
thermal waters, and other subjects involving the biology and
behavior of insects. He was not interested in economic entomol-
ogy and he believed that more intensive work on the biology of
insects was needed in view of the fact that comparatively little
is known even about many of our most common species. Im-
portant contributions were made by Lutz in the field of insect
behavior through his research work on insect reactions to ultra-
violet, on the training of bees to come to certain ultraviolet wave-
lengths and patterns for food, on his recordings and sound motion
pictures of insect sounds and insect behavior under various atmos-
pheric pressures. Such work required not only a thoughtful and
enquiring mind, an awareness of the pitfalls in conclusions, but
ingenuity in inventing mechanical devices needed for the tests.
These requirements Dr. Lutz possessed. In 1923 he was awarded
the Morrison Prize for his essay on ‘ ‘ The Colors of Flowers and
the Vision of Insects with Special Reference to Ultraviolet.” In
addition to his research work he did much to popularize ento-
mology and nature study. Tens of thousands of persons use his
4 ‘Field Book of Insects.” This was first published in 1918. A
second edition was published in 1921 and a third in 1935. Its
royalties put Dr. Lutz ’s four children through college.

In 1941 he wrote his last book entitled, “A Lot of Insects.”
This embodies accounts of the insects that, for the most part, were
the objects of Dr. Lutz’s curiosity, experimentation, and enter-
tainment over a period of many years, and includes his sound and
humorous entomological philosophy — all expressed interestingly
and in a lucid style. Dr. Lutz approached all his problems from
a stiumulating and thought-provoking viewpoint and this makes
for fascinating reading.

Dr. Lutz was a Fellow of the New York Academy of Sciences
and the American Association for the Advancement of Science,
a charter member of the Entomological Society of America and
its president in 1927, a member of the American Society of
Zoologists, the American Society of Naturalists, the Ecological
Society of America, Sigma Xi, Phi Beta Kappa, New York
Zoological Society, and the New York Entomological Society, his
presidency of the latter society having already been noted. He

Mar., 1944]

Weiss: Frank E. Lutz

was an advisor to the Buffalo Society of Natural Sciences and in
1937 a lecturer in Columbia University. He also served as chair-
man of the committee on Biological Relations Between Flowers
and Insects of the National Research Council.

Dr. Lutz made his home in New Jersey and funeral services
were held at his residence, 13 North Central Avenue, Ramsey,
New Jersey, on November 29, 1943. He is survived by his widow,
Mrs. Martha Ellen Brobson Lutz and four children, a son, Frank
Brobson Lutz, and three daughters, Anna Lutz, Ensign Laura
Lutz of the WAVES, and Mrs. Boyd Sherman.

These few paragraphs are but an inadequate summary of some
of the things accomplished by Dr. Lutz during his lifetime.
They fail to record many activites of which there are no records,
except in the memories of his friends. They fail to record the
happiness that Dr. Lutz’s chosen life-work brought to him, and
the pleasure that was his in creating problems and then solving
them. And they fail to mention his amiable and quizzical
philosophy of biological theories, flashes of which are apparent in
his writings. In a review of Dr. Lutz’s last book, and speaking of
his “Field Book of Insects” as well, Dr. H. M. Parshley said:
‘ ‘ They mirror a rare and admirable personality, a man of genuine
good will, a humorist, and one of a remarkable generation of
American naturalists.” — Harry B. Weiss.

Mar., 1944]

Bacon: Frank E. Lutz

BIBLIOGRAPHY OF FRANK E. LUTZ*

By Annette L. Bacon

1900. A study in the variations in the number of grooves upon the shells of
Pecten irradians (Lam.). Science, 12: 373.

1902. Report of Mr. Frank E. Lutz, Biologist. North Shore Improvement

Association. Reports on plans for the extermination of mosqui-
toes of the north shore of Long Island between Hempstead Harbor
and Cold Spring Harbor, p. 42-56.

The ecology of insect sounds. Can. Ent., 34: 64-66.

Inheritance of color among pointers. Science, 15: 571-572. (With
Eliz. B. Meek.)

1903. Note on the influence of change in sex on the intensity of heredity.

Biometrika, 2: 237-240.

Assortative mating in man, a cooperative study. Biometrika, 2 : 481—
498.

1904. Variation in bees. Bioh Bull., 6: 217-219.

1905. Biometry. Jour. Phil. Psych, and Sci. Meth., 2: 12-15.

Assortative mating in man. Science, 22: 249-250.

1906. Preserving spiders’ webs. Science, 23: 391.

The tegminal position in Gryllus. Can. Ent., 38 : 207-209.

1907. The merits of the fruit fly ( Drosophila ampelophila) . School Science

and Math., 7 : 672-673.

1908. Notes and literature: The effect of environment upon animals. Amer.

Nat., 42: 60-61.

The inheritance of the manner of clasping the hands. Amer. Nat.,
42: 195-196.

Notes on the inheritance of variations in the color pattern of Crioceris
asparagi. Psyche, 15 : 50-52.

The variation and correlations of certain taxonomic characters of
Gryllus. Carnegie Inst. Wash., Pub. No. 101, 63 p.

Combinations of alternative and blending inheritance. Science, 28:
317-318.

1909. Notes and literature: The effect of environment upon animals. Amer.

Nat., 43 : 55-57.

Notes and literature : The effect of environment upon animals. Amer.
Nat., 43: 248-251.

Experimental work with pomace flies. Amer. Mus. Jour., 9: 234-236.

1910. A brief history of antarctic exploration. A.M.N.H. Guide Leaflet 31,

27 p.

The annual scourge of flies and mosquitoes. Amer. Mus. Jour., 10:
183-185.

* Exclusive of book reviews and annual reports.

Journal New York Entomological Society

[Vol. LII

1911. Experiments with Drosophila ampelophila concerning evolution. Car-

negie Inst. Wash., Pub. No. 143, 40 p.

Relation between habit and structure in the insect world. Amer. Mus.
Jour., 11: 27-28.

Flea carriers of the plague. Amer. Mus. Jour., 11: 95-98.

Notes and literature: Mimicry. Amer. Nat., 45: 190-192.

Index to the generic names in Volume XXIX, Dr. Petrunkevitch ’s
1 1 Catalogue of Spiders of North, Central and South America and
Adjacent Islands.’ ’ Bull. A.M.N.H., 29:793-809.

1912. Inheritance of abnormal wing-venation in Drosophila. Proc. VII

International Zool. Congress (1907), p. 411-419. (Advance
reprint, 1910.)

String-figures from the Patomana Indians of British Guiana.

Anthrop. Papers A.M.N.H., 12: 1-14.

String-figures from the Upper Potaro. Timehri. Jour. Roy. Agric.

Com. Soc. British Guiana, 2(ser. 3): 117-127.

Do butterflies migrate? Amer. Mus. Jour., 12: 106-108.

The importance of insects. Amer. Mus. Jour., 12: 253—254.

1913. Experiments concerning the sexual difference in the wing length of

Drosophila ampelophila. Jour. Exper. Zool., 14: 267-273.

Eactors in aquatic environments. Jour. N. Y. Ent. Soc., 21: 1-4.

The distribution of Occidental spiders. Science, 37 : 567-568.

The life of the butterfly. House and Garden, August, p. 89-92.

1914. Humidity — a neglected factor in environmental work. Amer. Nat.,

48: 122-128.

Collecting in Cuba. Amer. Mus. Jour., 14: 99-106.

Biological notes concerning Drosophila ampelophila. Jour. N. Y. Ent.
Soc., 22: 134-138.

Our common butterflies. A.M.N.H. Guide Leaflet 38, 25 p. (With
F. E. Watson.) 4th ed., 1923, 31 p.; 5th ed., 1926, 21 p.; 7th
ed., 1941, 21 p.

How to collect and preserve insects. A.M.N.H. Guide Leaflet 39, 21 p.
3rd ed., 1917, 22 p.; 4th ed., 1920, 22 p.; 5th ed., 1924, 27 p.;
8th ed., 1936, 27 p.

1915. List of Greater Antillean spiders with notes on their distribution.

Annals N. Y. Acad. Sci., 26: 71-148.

Fragments of spider lore. Amer. Mus. Jour., 15: 424-426.
Experiments with Drosophila ampelophila concerning natural selec-
tion. Bull. A.M.N.H., 34: 605-624.

Outline plan of the hall of insect life. A.M.N.H. Circular, 4 p.

1916. Heredity and sex ; Mendelism and some of its recent developments.

Amer. Mus. Jour., 16: 228-242.

Faunal dispersal. Amer. Nat., 50: 374-384.

The geographic distribution of Bombidse (Hymenoptera) with notes
on certain species of Boreal America. Bull. A.M.N.H., 35: 501-
521.

Mar., 1944]

Bacon: Frank E. Lutz

Insects, an inexhaustible and relatively untouched field for recreation
or research. Amer. Mus. Jour., 16: 525—532.

1917. Heredity and sex. Mendelism and some of its recent developments

Scientific American Supplement, 84(2169) : 56-58.

1918. Field book of insects. G. P. Putnam’s Sons. New York, N. Y.

509 p.

General rules for the preparation of manuscript for the Bulletin and
Memoirs of the American Museum of Natural History. A.M.N.H.
Special Brochure No. 2, 14 p.

Insects and disease. A.M.N.H. Guide Leaflet 48, 73 p. (With C.-E.
A. Winslow.)

The control of insects injurious to gardens. A.M.N.H. Circular, 3 p.

1919. Scientific zoological publications of the American Museum for 1918.

Summary of work on invertebrates, fishes, amphibians, and birds.
Nat. Hist., 19 : 341-346.

Scientific zoological publications of the American Museum. Summary
of work on fossil mammals. Nat. Hist., 19: 731-733.

1920. Scientific zoological publications of the American Museum. Summary

of work on whales. Nat. Hist., 20: 107-109.

An entomologist in Colorado. Nat. Hist., 20: 312-325.

Our most brilliant butterflies. Country Life, 38(4): 27-33. (With
F. E. Watson).

Foreign insects newly come to America. Nat. Hist., 20: 501-502.
Notes on the distribution and bibliography of North American bees
of the families Apidse, Meliponidse, Bombidae, Euglossidae, and
Anthophoridae. Bull. A.M.N.H., 42: 491-641. (With T. D. A.
Cockerell.)

How crickets chirp. The Woodcraft Totem Board, 4(4) : 3.

How some insects walk on water. The Woodcraft Totem Board,
4(5): 3.

How moths and butterflies spend the winter. The Woodcraft Totem
Board, 4(6) : 3.

1921. How do insects make galls. The Woodcraft Totem Board, 4(7): 3.
How bees make honey. The Woodcraft Totem Board, 4(8) : 7.

How whales spout. The Woodcraft Totem Board, 4(9): 3.

How to collect insects. The Woodcraft Totem Board, 4(10) : 3.

How to preserve insects. The Woodcraft Totem Board, 4(11) : 3.
Geographic average, a suggested method for the study of distribution.
Amer. Mus. Novitates, 5, 7 p.

Field book of insects. Second edition. G. P. Putnam’s Sons. New
York, N. Y. 562 p.

1922. The reaction of Drosophila to ultraviolet. Science, 55: 519. (With

F. K. Richtmyer.)

Altitude in Colorado and geographical distribution. Bull. A.M.N.H.,
46: 335-366.

1923. Flowers and their insect visitors. Nat. Hist., 23 : 125-134.

Journal New York Entomological Society

[Vol. LII

1924. Apparently non-selective characters and combinations of characters,

including a study of ultraviolet in relation to the flower-visiting
habits of insects. Annals N. Y. Acad. Sci., 29 : 181-283.

Amateur entomologist and the Museum. A survey, from the half-
century mark, of the Department of Entomology, American
Museum. Nat. Hist., 24: 337-346.

Hunting stingless bees where east seems to be west. Nat. Hist., 24:
494-508.

Insect sounds. Bull. A.M.N.H., 50: 333-372.

1925. The new insect groups in the American Museum. Nat. Hist., 25:

126-135.

1926. Nature trails, an experiment in out-door education. A.M.N.H. Misc.

Pub. 21, 36 p. Second edition, 1931.

Taking nature lore to the public. Nat. Hist., 26: 111-123.

The friendly insects. Nat. Hist., 26: 147-151.

Principal orders of insects. Nat. Hist., 26: 164, with chart. (With
A. J. Mutchler.)

Our common butterflies. Nat. Hist., 26 : 165-183. (With E. E. Wat-
son.) (Reprint of Guide Leaflet 38.)

Insect sounds. Nat. Hist., 26: 206-213.

A nature trail in the sky. Nat. Hist., 26: 411-428.

1927. The still-open road. Nat. Hist., 27 : 373-382.

Wind and the direction of insect flight. Amer. Mus. Novitates 291,
4 p.

A much-abused but still cheerful cricket. Jour. N. Y. Ent. Soe., 35:
307-308.

1928. A new species of bacteria and the gall of an aphid. Amer. Mus. Novi-

tates 305, 4 p. (With E. Martin Brown.)

Little “beasts of prey” of the insect world. Nat. Hist., 28: 188-190.
Insects that erect tents. Nat. Hist., 28: 264-268.

1929. Experiments with “wonder creatures.” Nat. Hist., 29: 160-168.
Observations on leaf -cutting ants. Amer. Mus. Novitates 388, 21 p.

1930. An analysis by movietone of a cricket’s chirp ( Gryllus assimilis).

Amer. Mus. Novitates 420, 14 p. (With W. R. Hicks.)
Caddis-fly larvae as masons and builders. Nat. Hist., 30: 276-281.
Aquatic insect pets. Nat. Hist., 30: 389-401.

1931. Light as a factor in controlling the start of daily activity of a wren

and stingless bees. Amer. Mus. Novitates 468, 9 p.

Insects vs. the people. Nat. Hist., 31: 49-57.

In defense of insects. Sci. Mo., 32: 367-369. (Science Service Radio
Talks.) (Reprint of Insects vs. the people.)

Insects vs. the people. Condensed from The Scientific Monthly, April,
1931. Readers Digest, 19 : 359-360, 363.

In defense of insects. Article 28 in Science Today, edited by Watson
Davis, Harcourt Brace and Co., p. 184-188 (reprint).

Mar., 1944]

Bacon: Frank E. Lutz

A simple evaporimeter and some data obtained by its use in the Canal
Zone. Ecology, 12: 445-448.

Notes on the animal life of thermal waters in the Yellowstone National
Park. Amer. Mus. Novitates 498, 10 p.

1932. Our ignorance concerning insects. Can. Ent., 64 : 25-29, 49-56 and

73-78.

Experiments with Orthoptera concerning diurnal rhythm. Amer. Mus.
Novitates 550, 24 p.

1933. Experiments with 11 stingless bees” ( Trigona cressoni parastigma )

concerning their ability to distinguish ultraviolet patterns.
Amer. Mus. Novitates 641, 26 p.

li Invisible” colors of flowers and butterflies. Nat. Hist., 33: 565-
576.

1934. The il buckwheat problem” and the behavior of the honey bee.

Amer. Mus. Novitates, 688, 10 p.

Reactions of Drosophila to 2537 A radiation. Amer. Mus. Novitates
706, 14 p. (With E. N. Grisewood.)

World of the bee remains a mystery. New York Times Magazine,
Aug. 19, p. 12 and 15.

From low to high. Grand Canyon Nature Notes, 9 : 327-329.

1935. Field book of insects. Third edition. G. P. Putnam’s Sons. New

York, N. Y. 510 p.

1936. How about the tent caterpillar? Nat. Hist., 37: 149-158.

1937. Culture methods for invertebrate animals. 590 p. By P. S. Galtsoff,

F. E. Lutz, P. S. Welch, J. G. Needham, et al. Comstock Pub.

Co. Ithaca, N. Y. Collecting and rearing terrestrial and fresh-
water invertebrates, by Lutz, Needham, Welch, p. 40-50.

1938. The insect glee club at the microphone. Nat. Hist., 42: 338-345, 378.

1939. Big bees. School Nature League Bull. No. 10 (ser. 9), 3 p.

1941. The truth about termites. Nat. Hist., 48 : 113-115.

A lot of insects. G. P. Putnam’s Sons. New York, N. Y. 304 p.

1942. Insects, ticks and human diseases. A.M.N.I1. Guide Leaflet No. 113,

38 p. (With C. H. Curran.)

Mar., 1944]

Forbes: Lepidoptera

LEPIDOPTERA FROM WESTERN PERU AND
ECUADOR

By Wm. T. M. Forbes

Department of Entomology, Cornell University
Ithaca, New York

The occasion of this note is a little lot of Lepidoptera collected
by Mr. and Mrs. D. L. Frizzell in the arid northwest corner of
Pern, and Puna Id., Ecuador. While not many, the striking
character of the fauna is indicated by the presence of two new
Citheroniidae, and the region evidently is worth intensive collect-
ing. Among the normal and widespread things, may be men-
tioned Coea acheronta, Herse convolvuli from the Parinas Valley,
near Negritos, Peru; Pholus labruscce, Celerio annei, Utetheisa
ornatrix from the Parinas Valley; Hymenia fascialis, Eudioptis
hyalinata and Conchylodes arcifera from the Quebrada Mogollon.

The following are worthy of more specific mention. A female
Monarch shows the dull color and heavy black of the Lima
females. Single females of Ascia monuste from Negritos, Peru,
and Puna Id., Ecuador, suggest but hardly prove a racial dif-
ference.

Lycaena ramon Dognin. Parinas Valley, May 7, 1939. This
species was described from near Loja, Ecuador, a high temperate
and semiarid locality in the heart of the Andes1 but is equally
at home at sea level, where I took it commonly at Lima. It also
occurs on the western slope of the Andes at Chosica and Matu-
cana, and we have a specimen from Eten, a little north of Lima —
so it doubtless covers the whole arid area of western Ecuador
and at least northern Peru. We received the Eten specimen as
hanno, and other material may be floating around under that
name, but it is easily recognized by the ocelli on the hind wing
below — two larger between M3 and Cu2, and two only a little
smaller behind Cu2. The following key to the American species
of Lycazna, subgenus Hemiargus will place it more precisely.

1 Brown, Ann. Ent. Soc. Am., 34: 832.

Journal New York Entomological Society

[Vol. LII

1. Postmedial spots of fore wing below large and black, contrasting with the

small and fuscous subterminal series isola

-. Pm. spots of fore wing similar to subterminal ones ....; 2

2. Hind wing below with a large ocellus in cell M3 (cell 3 of Herrich-

Schaeffer system), similar to the one behind it 3

Hind wing with no ocellus in cell M3 5

3. Inner half of hind wing below contrastingly darkened, nearly obliterat-

ing the usual marks, which are much enlarged in this area; four small

ocelli . martha

Hind wing with ground all one color, the fuscous spotting generally
uniform :..... ......... 4

4. Two ocelli in anal area (behind Cu2) about two-thirds as large as the

ones in front of it and similar ramon

These ocelli small with a small silver spot only, or dull and similar to the
anterior subterminal markings :.... sachaeina

5. Anal area with a single large ocellus, similar to the one in front of Cu2 ... 6
Anal area with two subequal and inconspicuous spots or ocelli, about as

in zachaeina, or none .....I..:........... 8

6. Gray-brown below with spots all small, subequal and grayish ... bahamensis

-. Pale gray below a......... : :.. ." 7

7. Hind wing below, and above in female, with a very broad white submar-

ginal band ; all spots below small and similar dominica

-. Hind wing below with less conspicuous white submarginal band or none ;
three of the dark spots black and conspicuous ...... ammon ( catilina auct.)

8. Largely gray above; hind wing with slight tail and anal lobe bornoi

— . Mostly blue above; hind wing evenly rounded hanno ( catilina )

Goniurus jethira Butler. Easily recognized by the very large
honey-yellow spots, which are even larger in this specimen than
in the type. (Lep. Exot., p. 65, pi. 25, fig. 4). Parinas Valley,
May 7, 1939. The original locality was merely “Peru” but I
did not take it either at Lima or in the Chanchamayo, and suspect
it is a specialty of the arid Northwest.

Arsenura harrietse, new species

Closely similar to A. richardsoni Druce in major features; the fuscous
ground very lightly dotted with black, most definitely on posterior half of
median area. Antemedial of two widely separated blackish lines, the inner
nearly straight and outer much bowed out below cell, as in richardsoni, but
with heavier blackish filling, and the lines themselves more contrasting ; post-
medial line much further out, nearer to subterminal than to discal lunule,
blackish, more definite and more bowed out opposite lower angle of cell, the
median area conspicuously whitish toward inner margin; the following line
(subterminal in position, but probably morphologically the outer postmedian)
black, defined outwardly by a clay colored line, which is much finer and more

Mar., 1944]

Forbes: Lepidoptera

contrasting than the defining pale shade in richardsoni ; course much as in
richardsoni, but closer to margin, especially on costal third, and not nearly
as much extended toward base on inner margin; subterminal area not pale
as in richardsoni, but as dark as ground on fore wing and contrastingly
blackish on hind wing, with much paler yellow-brown terminal area. Sub-
apical black spot shorter than in richardsoni, not distinctly defined with
white; the terminal area below it vaguely shading between dull and red
brown, without the contrasting red-brown wedges of richardsoni. Discal
lunule black and contrasting with central tawny lunule, as in richardsoni,
but without the tawny bar extending from its outer side.

Hind wing generally similar to fore wing, but with only a faint darker
antemedial shade in place of the double line and dark filling, this shade
incorporating the faint discal bar (which is more distinct in richardsoni ) ;
postmedial band as on fore wing, much more conspicuous than in richardsoni ;
the outer pattern differing from the fore wing as noted.^ Wing form rounder
than in richardsoni, the apex of fore wing and angle of hind wing less
extended. Under side much less mottled than richardsoni, with three wavy
outer bands, varying from obsolescent to rather conspicuous, the outer
strongest and middle one weakest. Body plain brown as in richardsoni.

Expanse 92-110 mm., much smaller than richardsoni.

Puna Id., Ecuador; type and two paratypes in collection Cornell Uni-
versity.

This may possibly be a race of richardsoni, but the discontinu-
ous distribution, different wing form and many differences in
pattern suggest rather a good species. The following skeleton
key will place it in the genus :

1. Antemedial line double, of an outwardly oblique inner and a strongly

excurved outer element, the latter sometimes faint; st. space on pos-
terior half of fore wing and hind wing much broader than terminal

area I ...... 2

-. Am. line single, straight and outwardly oblique; both wings with a

sharply defined even slender pale marginal stripe *romulus

-. Am. line single, outcurved or angled, inwardly oblique to inner margin;
st. space narrower, usually much narrower than terminal space, except
sometimes for narrow extensions 4

2. Fore wing roundly falcate and deeply excavate below apex; hind wing

with tooth large, 12 mm. long on anterior side; hind wing with inner
st. line deeply sinuate, passing half way between margin and cell

at Mx ■„ *championi

-. Wings less irregular; st. line of hind wing crossing Mx two-thirds way
out to margin : 3

3. Discal spot of fore wing with a simple orange central lunule; margin of

wings hardly irregular *harrietae

Journal New York Entomological Society

t Vol. LII

Discal spot of fore wing with a short extension of middle of outer side
of lunule, forming a Greek e ; tail of hind wing 6 mm. long.

*ricliardsoni

4. Hind wing at least with a series of dark spots or lunules in terminal

area, wholly distinct from the subterminal complex; fore wing with
black markings conspicuous in cells Mx and almost always M2, the
upper usually joining to the apical pattern but conspicuous, the lower

usually free 5

-. Hind wing without this series of markings, though often with somewhat
similar extensions of the st. area; black patches in cells Mx and M2
normally absent, sometimes mere dashes, or lost in general blackish
shading : 11

5. Generally smaller species (female alcmene expanding 140 mm.), head

solid black, contrasting with the paler brown or fuscous thorax 6

-. Larger species (160 mm. and often more) ; head with at least a contrast-
ing pale bar over bases of antennae 8

6. Postmedial line not defined with pale; st. area expanded into a large

patch, occupying two-thirds the area between st. line and margin in

cells B4 and B5 . alcmene

-. Pm. line conspicuously defined by a following dirty white shading; st.
area below apex less extensive .4$;. 7

7. Markings of fore wing corresponding to the admarginal lunules of hind

wing taking the form of two similar large blotches in cells M3 and Cux

(feet not seen) pandora

-. This element of pattern taking the form of a waved diffuse admarginal

line ; tarsi concolorous dark brown *angulata

-. This element obsolete, except at anal angle, where it is not conspicuous;
tarsi cream white, contrasting *xanthopus

8. No black st. patch in cell M2; pm. area of both wings heavily shaded

with black; discal spot of fore wing lunulate; no admarginal spots on

fore wing sylla, hercules

-. Cell M2 heavily marked subterminally with black; discal spot a simple
bar . | : 9

9. No admarginal lunules on posterior part of fore wing; the black spot in

M2 fused with the blotch in cell M* aspasia

-. Spot in cell M2 separate, conspicuous, and followed with dark shades in
cells M3 and Cux similar to those on hind wing , 10

10. Ground with strong yellowish tint ; the two black patches in cells Mx and

M2 similar, very large, separated by hardly more than the light vein,

and scaled heavily with blue *meander

— . Ground with olive tint; the second black patch narrow and ovate.

*biundulata

11. Inner subterminal line nearly even, and marked with contrasting whitish

dots on veins or more irregular whitish patches; antemedial line when
distinct formed of a straight or concave bar across cell and a very
oblique lower portion from lower side of cell to basal angle 12

Mar., 1944]

Forbes: Lepidoptera

— . Inner st. line more irregular, normally with two large scallops between
M3 and Cu2, with only limited and irregular white marks; am. line
when distinct with lower portion less set off from portion crossing cell,
frequently in the form of a single excurved band 14

12. Lower segment of am. line conspicuous, in line with and more or less

continuing the black shade subterminally across cell Mx; a conspicuous

pale st. patch just below it in cell M2 ponder osa

— . Lower segment of am. line inconspicuous ; no single pale st. patch 13

13. Outer margin strongly irregular; postmedial area contrasting bright

chestnut brown loatesi

— . Margins less irregular; ground rather even dull light brown.

*crenulata

— . Margins still less irregular; pm. area somewhat contrasting, but light
brown arcaei

14. Discal spot lunulate, with contrasting pale center; dentations of inner

and outer st. lines if present not closely corresponding 15

— . Discal spots simple; the inner and outer st. lines closely parallel over
the dentations at cells M3 and Cux 16

15. Inner st. with distinct and outer with very strong dentations, the white

accompanying shade irregular *cymonia

— . Inner st. line nearly straight, and with even accompanying white shade;
outer st. obsolete thomsoni

16. Outer st. line with very strong, narrow black-filled dentations on both

wings, three or four of them on fore wing similar polyodonta

— . Outer st. line closely fitted to inner, both without strong dentations, save

for two in cells M3 and CUi of fore wing group *armida

— . Intermediate; the two upper teeth on fore wing present, but only half
as large as the two lower, and filled not with special black triangles
but with extensions of the generally blackish contrasting pm. area;
very large, expanding 175 mm * archianassa

In structural characters, the wing-form varies too widely in
obviously closely related species to serve for major subdivision,
but two groups are set apart by having pectinate antennae, sylla
and arcaei with their relatives — alternatives 5 and 12 of the key.
The residue, so far as seen, and including harrietce, have serrate
and fasciculate antennae.

Dysdwmonia species. There was badly broken material of a
very striking undescribed Dysdaemonia, with scalloped wings.
It will be described by Mr. Johnson, who has a better specimen
from the same region.

* Species represented in coll. Cornell University, many of them the gift of
Mr. Frank Johnson.

Journal New York Entomological Society

[Vol. LII

Givira tristani Schaus. A rubbed specimen from Puna Id. is
this species or very close.

Euclea copac Schaus. Puna Id. Agrees so far as can be seen,
but not good enough for certainty. The species was described
merely from “Peru.”

S 'eirocastnia elaphebolia Druce. Looks to me like a good spe-
cies. Described from Ecuador.

Monodes convexa, new species

Superficially similar to the North American festivoides group, but with
relatively much smaller body and arched fore wings. Structures of the nor-
mal Monodes, without sex-scaling ; male antennae ciliate, legs unmodified, with
a rough tuft beyond middle of mid tibiae; vestiture normal for Monodes, as
described by Hampson, but with the posterior thoracic tuft extended back,
almost completely covering the basal abdominal tuft, and roundly truncate
behind. Palpi with second joint upturned only a little beyond middle of
front, as also in normal species of Monodes.

Body fuscous, thorax somewhat mottled, with darker lower half of collar,
the upper half of front, vertex and lower half of collar contrasting blackish
in dark specimens; palpi with first and second joints with paler apices, the
outer sides contrasting blackish in dark specimens. Pore wing gray, varying
extremely in tint, light specimens with the terminal third much darker, dark
ones sometimes with the costal area rather darker. Costa with numerous
dark bars in light specimens, dark, cut with the pale gray ante- and post-
medial lines in dark specimens, and with about four small whitish bars
between postmedial and the pale apical shade. Subbasal line of vague paler
powdery scaling, toward costa, toward inner margin represented by an oblique
blackish bar; antemedial obscure, except for the contrasting pale bar at
costa; postmedial represented by dark spots on veins, followed by minute
whitish ones, broadly and evenly excurved on costal two thirds, then oblique
in to inner margin and slightly concave. Subterminal obscure, sometimes
indicated as the irregular boundary between a grayer terminal and browner
subterminal area; orbicular and reniform spots large, the orbicular outlined
with black except above, usually heavily before and behind; reniform very
large, only partly outlined; the area before orb. blackish, the filling between
orb. and ren. blackish or shaded with dark, without the sharp boundaries of
the festivoides group, the space between ren. and postmedial line usually
somewhat darkened, but without a blackish spot. Claviform minute, whitish,
usually contrasting, heavily outlined with black, especially before and beyond.
Apical oblique shade varying from obsolete to conspicuous, cream white to
ash gray, with the basal half shaded with buff in light specimens and slightly
warmer brown in dark ones; starting from pm. line opposite cell, curving up
and widening in a horn shape, and ending on outer tenth of costa; partly
edged behind with black. Terminal blackish dots, obscure in dark speci-

Mar., 1944]

Forbes: Lepidoptera

mens, preceded by whitish points, alternating with the pm. ones. Fringe
powdery gray, the outer half rather smoother and less powdery. Hind wing
dirty white, shading into fuscous on outer half; alula cream, clothed with
large scales and conspicuous. Expanse 17-20 mm.

This species in normal specimens will run in Hampson’s key to
the festivoides group, from which it is distinguished by the total
lack of warm coloring, much smaller body and arched fore wings,
also in maculate forms by the less sharply defined blackish about
the orbicular and reniform. It is probably close to bogotana
Felder and aphaidropa Dyar (which are presumably merely color
forms of each other) but is smaller, and none of our series are as
pale as Felder’s figure of bogotana, nor show the reddish shadings
along costa of aphaidropa. It varies enormously, from speci-
mens (males) with the basal two thirds luteous and only the outer
third blackish, much like Felder’s figure, to specimens (females)
that are wholly suffused with smoky gray and coal black, with
all the markings obsolescent. Such specimens will probably key
out to phceopera, but differ from it and its relatives by the rather
even dull gray, with the darker area between orbicular and reni-
form, and the paler apical area at least faintly visible.

The male genitalia of Monodes are extremely varied, and con-
vexa resembles nucicolora, grata or fusimacula more closely than
festivoides, having the valves slender, with a complicated basal
chitinization composed of parts of sacculus, costa, and perhaps
clasper, and sending a spike forward across the costa; weak
clasper at a third way out, crossing costa; juxta slenderly ex-
tended as a complete anellus, and transtilla also bent into a round
arch closely parallel to it. MCdoeagus with two massive spines,
formed of fused cornuti.

Holotype, male from Chosica, Peru, May 25, 1920. Numerous
paratypes of both sexes, from Chosica and Lima, Peru, May
1920; a couple caught by Parish at Lima in 1915; and a pair
from the Amotape Mts., N.W. Peru, collected by the Frizzells;
all in collection Cornell University. I believe other specimens
exist in collections, perhaps labelled bogotana, but the Lima
fauna has been extraordinarily neglected, considering how many
good collectors have passed through it or even used it for a base.

Cydosia phcedra Druee. Puna Id.

Cobubatha numa Druce. Amotape Mts.

Journal New York Entomological Society

[Vol. LII

Dichochroma, new genus

Similar to the Pyraustine genus Dichogama. Yestiture of large, smooth
scaling; palpi upturned to middle of front, close-scaled, slightly flattened
against the front; the segments well marked off, third segment larger than
in Dichogama, two-thirds as long as second, continuing the direction of sec-
ond; maxillary palpi rough-scaled, flattened against the face, as in Dicho-
gama. Tongue strong. Legs very short for a Pyraustine, as in Dichogama,
mid tibia about as long as femur without trochanter, and tarsus hardly
longer. Fore wing presumably with the long Arctiid-like frenulum hook of
Dichogama (male not seen) ; R3 and R4 stalked, R5 stalked with M4, well
separated from R4, M2 and M3 stalked, Cu4 parallel to M3, arising well before
end of cell. Hind wing without fringe on Cu; M2 and 3 strongly stalked, as
in D. fernaldi, but unlike the other Dichogamas at hand.

Third A of fore wing is strong and makes a wide loop, but I cannot see
if it runs back into 2d A.

This genus is clearly a development of Dichogama, differing
from it, and from all Pyraustinae known to me in the stalked
R5 and Mi (veins 6 and 7) and from most in the stalking of
M2 and M3 in both wings. It is also far smaller than any Dicho-
gama yet known, and is found on the Mainland, while Dichogama
is essentially Antillean, only D. diffusalis not yet being known
from the Antilles. In Hampson’s key (Proc. Zool. Soc., 1898,
594) it will run to Hymenia or Macaraetera, according to how the
third segment of the palpus is interpreted, but has no real kin-
ship to either. The stalked veins will easily separate it.

By the way there is no reason for marking several species of
Dichogama “incertae sedis” as Klima does in Lep. Cat. 89, p. 122.
We have colotha, fernaldi and gudmanni from Porto Rico and
they are normal Dichogamas, save for the stalked instead of ap-
proximate M2 and M3 in fernaldi ; and amalyilis and bergii show
patterns that would hardly occur elsewhere ; but smithii, unques-
tioned by Klima, is the well known Noctuid, Casandria abseuzalis,
which I found common in Porto Rico.

Dichochroma muralis, new species

Head and thorax mouse gray, dusted with whitish scale-tips; thorax im-
maculate ; shaft of antennae blackish ; palpi with first segment whitish, second
mouse gray, but whitish along the ventral inner keel; third segment blackish
with contrasting dirty white tip. Under side cream white, the front side of
the fore legs fuscous. Abdomen above mouse gray, immaculate, below
nearly white.

Mae., 1944]

Forbes: Lepidoptera

Fore wing mouse gray, immaculate but with pale scale-tips especially
toward base, and sparsely overlaid with narrow whitish strap-shaped scales;
fringe whitish. Hind wing translucent white with narrow and broken fus-
cous terminal line. Expanse 16 mm.

Amotape Mts., N. W. Peru, H. & D. L. Frizzell; type one female in coll.
Cornell University.

In sum this little fauna from the north end of the arid coastal
strip of South America is a curious one. While the typical mate-
rial of this strip is present ( e.g ., Celerio annei and Monodes con -
vexa) there is also a definite Central American element (notably
the Arsennra, which is closest to A. richardsoni) , and a few spe-
cies which now appear to be endemic, such as the undescribed
Dysdsemonia, Euclea copac and Dichochroma murina. Plainly
many more interesting things are due to come out of the area.

Mar., 1944]

Miller: Drosophila

DROSOPHILA MELANURA, A NEW SPECIES OF THE

MELANICA GROUP

By Dwight D. Miller*

California Institute of Technology, Pasadena, California

During most of the summer of 1941 the author collected Droso-
philas and related forms on the River Campus of the University
of Rochester at Rochester, New York. One of the commonest
types to be found in the fermented banana traps used in the col-
lections was a form identified as Drosophila melanica Sturtevant
(1916). From June 5 through August 25 there were gotten
1263 individuals of this species, 538 of which were females, 725
males. In a recent paper Patterson (1942) has indicated that
D. melanica should be divided into two subspecies, melanica and
paramelanica, differing in distribution, melanica having been
found in the southeastern United States and in Mexico, para-
melanica occurring in the northeastern part of this country. It
is probable, on the basis of locality, that the D. melanica indi-
viduals collected at Rochester belonged to the subspecies para-
melanica. Towards the end of the collecting period, from
August 19 through August 25, there occurred in the traps some
male Drosophilas that seemed to differ from the D. melanica
males only in the unusually dark pigmentation of their genital
region. Altogether there were 10 such males collected. Since
it was suspected that these males represented a new form, a num-
ber of apparently D. melanica females collected at the time were
isolated individually into culture bottles in the hope that some
of them would have offspring, males of which would be of this
new type. Unfortunately, none of the females so isolated bred.
Since it seemed unlikely that the females would have any progeny
if left to themselves, an attempt was made to mate them to some
of the melanica- like males. Culture bottles that had contained
one of the females with two such males yielded some offspring.
The male progeny were all of the new type in that their genital

* This study was begun while the author was holding a teaching assistant-
ship at the University of Rochester, Rochester, New York.

Journal New York Entomological Society

t Vol. LII

region was darkly pigmented. These offspring proved to be fer-
tile, and through allowing them to mate among themselves a
strain was derived. Males of succeeding generations have all
persisted in differing from D. melanica males in the dark pig-
mentation of the genital region. On the basis of this constant
character of difference, as well as because of others noted below,
the melanica- like form collected at Rochester is here designated
a separate species, Drosophila melanura. A description of the
new species is given next, with notes on comparison of the new
form with D. melanica following just afterwards.

Description of the Species

Drosophila melanura, sp. nov.

External Characters of Imagines.

$ Arista with about 9 branches. Antennas brownish, pollinose. Front
blackish. Middle orbital length about i that of posterior one. Second oral
length about ^ that of first. Carina broad below, sulcate. Face brownish.
Cheeks pale brownish, about i greatest diameter of eye at their greatest
width. Eyes red, with black pile.

Acrostichal hairs in irregular rows; about six rows at the anterior dorso-
central bristles. Anterior scutellars convergent. Mesonotum dull brown,
with four slightly paler longitudinal stripes, two extending forward from
just inside anterior dorsocentral bristles, two lying just outside the lateral
pairs of dorsocentral bristles. Pleurae brownish. Sterno-index about 0.9.
Legs pale brownish. Two apical bristles on first tibiae, one on second; pre-
apicals on all three.

Broad, brown bands on abdominal segments 2 through 6, with posteriorly
narrowing median interruptions, with but little interruption near the lateral
margins of the tergites. Genital region (genital arch, anal plates, claspers,
penis apparatus) dark brown.

Wings clear, veins brown. Costal index about 2.9; 4th vein index about
1.4; 5x index about 0.9; 4c index about 0.6. Two bristles at apex of first
costal section. Third costal section with bristles on its basal

Body length about 3.3 mm. (alive) ; wings 2.8 mm.

$ Broad, brown bands on abdominal segments 2 through 6, with posteri-
orly narrowing median interruptions, often with wide lateral interruptions
on the 6th segment.

Body length about 3.5 mm. (alive) ; wings 3.0 mm.

Internal Characters of Imagines.

Testis with about 3 inner coils (probably the seminal vesicle) and 3 outer
ones, the coiled portion tending to be orange. Ejaculatory sac with 4 long
diverticula, 2 anterior and 2 posterior.

Spermathecae chitinized. Ventral receptacle with about 30 coils.

Mar., 1944]

Miller: Drosophila

Additional notes.

Egg. — With 2 filaments, each about f the length of the egg.

Puparium. — Amber. About 9 branches in each anterior
spiracle. Horn index (length of puparium/length of anterior
spiracle horn) about 12.

Chromosomes. — Female metaphase plate contains : one pair of
large V’s, two pairs of rods, one pair of medium V’s, and one pair
of small V’s. Male metaphase plate has a J in place of one of
the large V ’s.

Key Position. — The following is to be taken as an expansion
of the first line of couplet 50 of the Key to North American Spe-
cies of Drosophila of Sturtevant (1942), to which point an
attempt to classify a specimen of D. melanura should lead one.

50a. Male genital region dark brown melanura

Male genital region not dark brown melanica

Distribution. — This species has been collected only in a small
wood on the campus of the University of Rochester (River
Campus), Rochester, New York.

Type specimens of D. melanura are deposited in the American
Museum of Natural History, New York, N. Y.

Comparison with D. melanica

Drosophila melanura has been compared with individuals
taken from D. melanica strains kept in this laboratory. These
strains have been classified as to subspecies ( melanica or para-
melanica) by Prof. A. H. Sturtevant. The above description of
D. melanura , as well as the notes which follow, is based largely
on individuals of the single existing strain of this species, de-
rived as indicated in the first paragraph.

The impression has been gotten that D. melanura is a larger
form than D. melanica. This has been gotten not so much from
wild individuals as from flies raised in the laboratory (the size
measurements given in the above description were taken from
“average” laboratory individuals). The difference in size may
be partly due to the fact that the new species has not bred as
vigorously in the laboratory as has D. melanica and has usually
had the advantage of relatively uncrowded culture bottles.

Journal New York Entomological Society

[Vol. LII

D. melanura has also seemed to have a lighter body color than
D. melanica. This difference is especially noticeable on the dor-
sal surface of the thorax, which may be described as medium
brown in the new form, dark brown or very dark brown in D.
melanica. Accompanying this difference is a greater prominence
of the mesonotal stripes in melanura than in D. melanica. It is
the impression of the author that the D. melanica subspecies tend
to differ from each other in that melanica is somewhat darker
than paramelanica. The thorax color difference between me-
lanura and D. melanica melanica has been quite striking. Since
a body color difference was not appreciated at the time the col-
lections of melanura and D. melanica (probably paramelanica)
were being made, the separation of wild females on the basis of
color was not attempted. It remains to be seen how reliable a
criterion this is for the identification of wild individuals in
general.

The abdominal banding pattern of D. melanura males was
found to be different from that of the D. melanica males exam-
ined. Whereas in melanura males the band on the 6th segment
(as well as the others) was always found to be complete laterally,
this was never seen to be the case in D. melanica males. In a
D. melanica strain ( paramelanica ) from South Amherst, Massa-
chusetts, the males’ abdominal bands were all found to become
indistinct near the lateral margins of the tergites. In a D.
melanica strain ( melanica ) from Walnut Creek, Texas, while the
bands on the 2nd through 5th abdominal segments generally
extended all the way to the edges to the tergites, the band on the
6th segment was interrupted laterally.

Mention has already been made of the darkly pigmented geni-
tal region of males of the new species. This seems to be the best
character whereby it may be recognized as different from D. me-
lanica. In D. melanura males the genital arch, anal plates, and
claspers become dark brown a few days after emergence. The
plates of the penis apparatus also become dark. The result is
that the genital region stands out conspicuously against the light
ventral abdominal wall. In D. melanica the male genital region
seems relatively inconspicuous.

In D. melanura and in D. melanica the penis apparatus appar-
ently consists of two rather broad lateral plates as well as the

Mar., 1944]

Miller: Drosophila

structure probably homologous to the chitinous rod called the
penis by some authors (for example, Nonidez, 1920, in D. melano-
g aster). The appearance of the lateral plates was found to

Fig. 1. A. Male genital region of B. melanura. B. Male genital region
of B. melanica paramelanica (South Amherst, Mass.). The labels indicate:
the anal plates (a), claspers (c), genital arch (g), and penis apparatus (p).
These drawings were made from dead specimens, and the penis apparatus is
shown farther forward than its usual position in living, etherized individ-
uals. C. Penis apparatus of B. melanura. D. Penis apparatus of B.

melanica melanica (Walnut Creek, Texas). E. Penis apparatus of B.
melanica paramelanica (South Amherst). C, D, and E are side view draw-
ings made on a somewhat larger scale than A and B. The edges labelled
with a “p” in C and E correspond to the edges so labelled in A and B
respectively. The point labelled with an “x” in D was found to vary some-
what in prominence.

differ between melanura and D. melanica. In melanura they
were deeply incised at the anterior edge (Figure 1, C), whereas

Journal New York Entomological Society

[Vol. LII

in D. melanica they were not so much or scarcely at all so incised
(Fig. 1, D & E). The structure of the plates in D. melanica
melanica strains examined (Walnut Creek, Texas; Coffeeville,
Kansas; Taneycomo, Missouri) seemed intermediate (Fig. 1,
D) between that of D. melanura (Fig. 1, C) and that of D. me-
lanica paramelanica strains (Madison, Wisconsin; South Am-
herst, Massachusetts) (Fig. 1, E). In D. melanura the impres-
sion was gotten that the penis apparatus is somewhat larger and
more conspicuous relative to its surroundings than in D. melanica
(Fig. 1,A&B).

The anterior spiracle horns of the puparium were found to be
relatively shorter in D. melanura than in D. melanica. The horn
index (length of puparium/length of horn) was determined to
be about 12 in melanura, whereas melanica estimates were 9 for
the South Amherst strain ( paramelanica ) and 7 for the one from
Walnut Creek {melanica).

The chromosomes of D. melanura were investigated by means
of acetic orcein smear preparations of larval ganglia and of
ovaries and testes taken from pupse and adults (using the 70 per
cent acetic acid stain of LaCour, 1941). Examples of chromo-
some groups are given in Figure 2, A, B, and C. It may be seen
that the smallest chromosome is not dot-shaped, as was reported in
D. melanica by Metz (1916), but, rather, a small V. An investi-
gation of D. melanica chromosomes, both paramelanica (South
Amherst and Madison) and melanica (Walnut Creek), has con-
firmed Metz’ report of dot-like chromosomes in this species
(Figure 2, D, E, and F). Of interest here is the statement of
Griff en (1942) that the melanica group species D. nigromelanica
Patterson and Wheeler (1942) has rod-shaped microchromosomes
rather than dots. The salivary gland chromosomes of D. me-
lanura have not been studied.

As may be seen from the first paragraph of this paper D.
melanura males were gotten in the summer, 1941, collections at
Rochester in a much smaller number than were D. melanica
males. Moreover, D. melanura was only collected past the
middle of summer, late in August, while D. melanica occurred in
the traps by early June. An ecological difference between the
two species is suggested. D. nigromelanica was collected also

Mar., 1944]

Miller: Drosophila

and had an occurrence in the traps similar to that of D. me-
lanura; from July 26 to August 25 there were gotten 26 indi-
viduals of this species, 11 of which were males. A record of the
collections of these melanica group species is given in Table 1.

Fig. 2. A and B. D. melanura ovary metaphase figures. C. D. melanura
testis metaphase. D and E. D. melanica paramelanica (South Amherst)
ovary metaphases. In E some of the chromosomes, including the dot-like
ones, are obviously split. F. D. melanica melanica (Walnut Creek) testis
metaphase. The dots are close together. G, H, and I. Metaphase figures
found in the testes of hybrids derived from melanica (Walnut Creek) females
mated to melanura males.

D. melanura has been kept with some difficulty on the Droso-
phila culture media in use at this laboratory. The impression
has been gotten that D. melanica has not been so difficult to main-

Journal New York Entomological Society

[VOL. LII

tain. In December, 1941, onr corn meal-molasses-agar medium
was changed in that agar was omitted from it and rolled oats
were added (formula of Dr. R. H. MacKnight). The result has
been a somewhat softer, wetter medium. From soon after the

Table 1. Week-by-week record of melanica group species collections at
Rochester, New York, during the summer of 1941. The number of D.
melanura females gotten is not recorded; these were most probably all classi-
fied as D. melanica females. Since the circumstances of collection (number
of traps, number of collections, etc.) varied from week to week, the variation
in absolute numbers should not be taken to reflect very well the changing
state of the wild population.

Collections for the week beginning: .

June 1 June 8

June 15

June 22

June 29

melanica

$ $

nigromelanica

melanura

July 6

July 13

July 20

July 27

Aug. 3

melanica

$ $

171

134

115

$ 2

111

125

nigromelanica

$ $

melanura

Aug. 10

Aug. 17

Aug. 24

Total

melanica

$ $

725

$ $

538

1263

nigromelanica

$ $

melanura

$ $

change was made until about the middle of May, 1942, the D.
melanura strain refused to breed at all. Following this period
there has been a gradual recovery of ease of culture.

Hybrids with D. melanica

A few attempts were made to cross D. melanica females with
D. melanura males, several individuals of a kind being used in
each mating. Progeny have been gotten both from crosses of
melanica (Walnut Creek) females by melanura males and from
paramelanica (Madison) females by melanura males (Table 2).

Mar., 1944]

Miller: Drosophila

A few matings were also made between D. nigromelanica females
and melanura males, using a Harden County, Texas, strain fur-
nished by Prof. J. T. Patterson, but no progeny were gotten
(Table 2).

In general the interspecific cross progeny have been not very
different from the maternal species. The penis apparatus of the

Table 2. Interspecific crosses involving D. melanura. The flies were
put into fresh culture bottles on the dates listed under each mating. The
numbers of progeny are placed opposite the dates appropriate to the bottles
in which they occurred.

Cross

Progeny

$ $ $ $ Totals

D. melanica melanica $ $ xD. melanura $ $

Walnut Creek

$ $ x Rochester $ $

(7 of each)

8- 7-42

8-10-42

8-15-42

Walnut Creek

$ 9 x Rochester $ $

(22 of each)

8- 8-42

8-11-42

8-15-42

Walnut Creek

$ $ x Rochester $ $

(15 of each)

8-10-42

8-15-42

8-19-42

8-24-42

8-31-42

9- 8-42

137

111

248

D. melanica paramelanica $ $ xfi. melanura $ $
Madison $ $ x Rochester $ $ (5 of each)

8- 8-42
8-11-42
8-15-42

Madison $ $ x Rochester $ $ (7 of each)
8-10-42
8-15-42
8-19-42

Journal New York Entomological Society

[VOL. LII

Table 2 — ( Continued )

Cross

D. melanica paramelanica $ $ x D. melanura $ $
(cont.)

Madison 9 $ x Rochester $ $ (15 of each)
8- 7-42
8-10-42
8-15-42
8-24-42

8- 31-42

9- 8-42

D. nigromelanica $ $ xD. melanura $ $

Harden Co. $ $ x Rochester $ $ (8 of each)
9- 8-42
9-12-42

Harden Co. $ $ x Rochester $ $ (4 of each)
9-1-42
9-4-42
9-8-42

Harden Co. $ $ x Rochester $ $ (13 of each)
9- 4-42
9- 8-42
9-10-42

2 $

Progeny
$ $ Totals

27 56

hybrid males, for example, has been rather similar to that of
males of the form to which the mother belonged. Both sexes
have seemed quite viable, and the sex ratio has been about 1 to
1 (Table 2).

Metaphase plates found in the testes of hybrid males derived
from both the kinds of crosses mentioned above contained one
dot-like chromosome and one small V (Fig. 2, G, H, and I). This
evidence may be offered as one form of proof of the hybrid nature
of these offspring. The salivary gland chromosomes of several
larvae were looked at, and although a very careful study of these
chromosomes was not made, it did seem likely that they were
heterozygous for a number of chromosomal rearrangements,
probably inversions.

Mar., 1944]

Miller: Drosophila

Table 3. Backcrosses of hybrid females to B. melanura and to D.
melanica.

Cross

Progeny

$ $ $ $ Totals

( melanica 9 2 x melanura) $ 9 x B. melanura $ $

(W. C. $ $ xRoch.) $ 9 x Rochester $ $ (13
of each)

8- 31-42

9- 3-42
9- 8-42

(W. C. 9 $ xRoch.) 9 $ x Rochester $ $ (7
of each)

9-1-42

9-4-42

9-8-42

( paramelanica 9 2 x melanura ) 9 9 xD.
melanura $ $

(Mad. 9 2 xRoch.) 9 $ x Rochester $ $

(2 of each)

9- 4-42
9- 8-42
9-10-42

( melanica 9 $ x melanura) 9 $ x B. melanica
melanica $ $

(W. C. 9 $ xRoch.) 9 2 X Walnut Creek $ $
(17 of each)

9- 4-42

9- 8-42

9-10-42

9-18-42

9-25-42

102

(. paramelanica 9 9 x melanura) 9 9 x para-
melanica $ $

(Mad. 9 2 xRoch.) 9 9 x Madison $ $
(9 of each)

9-30-42

10- 3-42
10- 8-42

Journal New York Entomological Society

[Vol. LII

No offspring have been gotten from hybrid males and females
kept together. It seems likely that the males are sterile.
Although hybrid testes were found to contain many sperms, no
motility of these sperms in Kingers’ solution was seen. More-
over, acetic orcein smears have shown these testes to contain a
number of rather large, spindle-shaped, dully-staining bodies,
but no regular sperm heads. It is probable that these elongate
structures are the product of an abnormal spermiogenesis.

Attempts have been made to backcross hybrid females both to
D. melanura and to D. melanica males, and offspring were gotten
in the case of a group mating of hybrid females (Walnut Creek
22 x Rochester to Walnut Creek males ( melanica ) (Table
3). A rather obvious excess of females over males may be seen
in these progeny. Since no mutants were involved in the crosses
that gave rise to them, little could be concluded about the genetic
constitution of the back-cross individuals. Some of the males
were dissected 10 or 11 days after emergence and their testes
examined. Out of 20 such males 2 had testes containing small,
pyknotic bodies of various shapes, but no normal sperm heads,
while the others had normal appearing sperms. Several recently
hatched males were dissected and their testes examined for
chromosome groups. In one of the testes there were found
rather clear figures showing the presence of a dot and small Y
together, and in this testis there were a number of quite normal
appearing sperm heads.

SUMMARY

Drosophila melanura, a new species similar to D. melanica
Sturt., is described here. Notes on comparison of the two forms
are presented. These species differ cytologically in the appear-
ance of the smallest chromosome in the metaphase plate, D. me-
lanura having a small V where D. melanica has a dot. Hybrids
have been obtained from crosses between D. melanura females
and D. melanica males, and the female hybrids have proved to be
fertile.

ACKNOWLEDGMENTS

The author wishes to thank Dr. H. D. Stalker of Washington
University, St. Louis, Missouri, and Prof. A. H. Sturtevant of

Mar., 1944]

Miller: Drosophila

the California Institute of Technology for advice and assistance

in connection with this study.

LITERATURE CITED

Griffen, A. B. 1942. Relationships in the Melaniea Species Group. Univ.
of Texas Publ., 4228: 68-73.

LaCour, L. 1941. Acetic-orcein: A New Stain-fixative for Chromosomes.
Stain Techn., 16: 169-174.

Metz, C. W. 1916. Chromosome Studies on the Diptera. III. Additional
Types of Chromosome Groups in the Drosophilidse. Amer. Nat., 50 :
587-599.

Nonidez, J. F. 1920. The Internal Phenomena of Reproduction in Dro-
sophila. Biol. Bull., 39 : 207-230.

Patterson, J. T. 1942. Interspecific Hybridization in the Genus Drosoph-
ila. Univ. of Texas Publ., 4228: 7-15.

and M. R. Wheeler. 1942. Description of New Species of the

Subgenera Hirtodrosophila and Drosophila. Univ. of Texas Publ.,
4213: 67-109.

Sturtevant, A. H. 1916. Notes on North American Drosophilidse with
Descriptions of Twenty-three New Species. Ann. Ent. Soc. Amer.,
9: 323-343.

. 1942. The Classification of the Genus Drosophila, with Descrip-
tions of Nine New Species. Univ. of Texas Publ., 4213 : 5-51.

Mar., 1944]

Brown: Pieridje

NOTES ON MEXICAN BUTTERFLIES, II, PIERID^

By F. Martin Brown

The collections made by Hoogstraal’s parties and by Potts con-
tained 33 of the 71 species of Pieridse reported by Hoffmann.
Those lacking are primarily the truly tropical species that enter
Chiapas and the southern lowlands and some of the western
Nearctic things that are found in Sonora and Baja California.
I have added the data from a small collection made by H. D.
Thomas in 1936.

Several localities not mentioned in the first paper on the
Papilionidae (g.u.)# are noted here:

Geographic Data

Acahuato, Michoacan, 19° 20' N., 102° 20' W., 3000 ft.

“On the edge of the plateau above Apatzingan. Open semi-
desert with scattered trees below the town and open pine forest
above it. Transitional between the tropics and temperate area. ’ ’
Hoogstraql.

Cumbres, Vera Cruz, 6000 ft., km. 295 on road to Vera Cruz
from Mexico City.

“North and west of Orizaba, very definitely up and out of the
tropical zone and rain belt, into a dry upland type of country
even though still the bottom of the valley. A lot of straggling
thorn bush, tremendous organ cactus, a lot of Opuntia. Short
grass and rock slopes. Temp, at about 75 or less.” Potts.

El Mante, San Luis Potosi.

“A strange conglomeration in this region of swampy country
with tall palms and very tall thorn bush jungle next to bananas,
corn fields and maguey. Cactus growing in mud ! Temp, proba-
bly 95° F.” Potts.

Hda. Potrero Viejo, nr. Paraje Nuevo, Vera Cruz.

‘ ‘ Cultivated country for centuries, yet plenty of original vege-
tation nearby. I collected only in a small meadow near the
* Jour. N. Y. Ent. Soc., Yol. 51, No. 3, p. 161-178, 1943.

100

Journal New York Entomological Society

[Vol. LII

hacienda itself.” Potts. Between Fortin and Orizaba on the
Mexico City-Vera Cruz highway.

Hda. Santa Engracia, Tamaulipas.

I have no data on this station. F.M.B.

Monte Alban, Oaxaca, 16° 50' N., 96° 20' W., 6000 ft.

“ Hills back of Oaxaca, the only moist country nearby!”
Potts.

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 26° 40' N., 100° 15'
W., 1000-1600 ft.

‘ ‘ Collections were made up to 1600 ft., low shrubs and cactus.”
Hoogstraal.

Orizaba, Vera Cruz, 18° 45' N., 96° 50' W., 2200 ft.

“About a kilometer SE. of the town. Just the top end of the
tropics or at least the jungle tropics, along the valley slope.
Impenetrable underbrush, lush, rains some 9 or 10 months of the
year here. ” Potts.

Sta. Lucrecia.

A Thomas station which must be on the west coast, probably in
Michoacan, from the material so labeled. F.M.B.

Rio Balsas, Guerrero, 18° dz S., 98°-102° W.

The great river basin of southern Mexico. ‘ ‘ The river itself is
pretty dry. Collected up a side canyon to the south. Temp.
105-110, muggy. Thorn bush and rocks.” Potts. The collec-
tion was made near Mexcala (18° 0' S., 99° 30' W., 2400 ft.).

Tuxpango, Vera Cruz, ? 1500 ft.

“Top of a big wide barranca SE. of Orizaba. County simi-
lar to ‘Orizaba.’ ” Potts.

Dismorphiinae

I am surprised that these two collectors, Hoogstraal and Potts,
were so unsuccessful in finding members of this sub-family. At
least nine species are known from the areas they visited. Only
Hoogstraal collected any and his material represents one species.

Mar., 1944]

Brown: Pierid^:

101

61. Enantia jethys Boisduval.

G. & S. (1), 2: p. 182-183, 727, pi. 61, f. 56; pi. 74, f. 27,

28 (as albania).

R. (2), p. 100.

H. (3), p. 654-5.

Tancitaro, Michoacan, 6600 ft. 3 J'.J' 2 52 vii.20-viii. 14.40
(H.H.).

Hoffmann, p. 654U5, recognizes three species, citronella Felder
(1861), jethys Bdv. (1836) and mita Reak (1866). I feel that
these represent but one variable species. I have a long series of
specimens from various tropical stations and among them are all
intergrades from jethys to citronella to mita. Whether these are
in turn only varieties of melite Linnaeus or not I am not sure.
The most heavily marked specimen should be referred to f. jethys
Bdv., those without the dark bar along the inner margin of the
forewing to f. citronella Felder, and those with further restric-
tion of the dark apical markings to f . mita.

The specimens noted above are of f. citronella. I suspect that
the degree to which the dark markings are developed is related
to the season or moisture, the darkly marked individual being
1 1 wet season ’ ’ the lightly marked, 1 1 dry season. ’ ’

63. Enantia marion, Godman & Salvin.

G. & S., 2 : 184.

R., p. 100.

H. , p. 655.

El Sabino, Uruapan, Michoacan, 1 vii. 15-30.36
(H.D.T.).

Pierinas

(Euchloini)

72. Hesperocharis costaricensis Bates.

G. & S., 2 : 126, pi. 60, f. 3, 4.

R., p. 79, pi. 23f.

H, p. 656.

Rio Blanco, Yera Cruz, 2200 ft., 1 J1 v.10.41 (R.P.).

El Sabino, Uruapan, Michoacan, 1 2 vii. 15-30.36
(H.D.T.).

102

Journal New York Entomological Society

[Vol. LI1

Apparently this is the first record of the species from the east
coast of Mexico. Hoffmann, p. 656, reports the species from the
west coast of Guerrero, Michoacan, Jalisco and Colima.

(Rhodocerini)

74. Colias eurytheme Boisduval.

G. & S., 2 : 151 (as chrysotheme) .

R., p. 93, pi. 271

H. , p. 656.

Hda. Yista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 2 ?5 vi.17.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 J1 vii.30.39 (H.H.).
Tancitaro, Michoacan, 6600 ft., 1 £ viii.11.40 (H.H.).
These few specimens do not indicate the abundance of the
species in Mexico. It is probable that the species was ignored in
most localities. The Nuevo Leon material is f. amphidusa Bdv.,
the other is typical of f . eurytheme .

77. Zerene caesonia Stoll.

G. & S., 2 : 153, 727 (as cesonia).

R., p. 94, pi. 261

H. , p. 656.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 4 1 $ vi.16-18.40 (H.H.).

Arroyo del Meco, Tamaulipas, 1320 ft., 1 $ iv.28.41

(R.P.).

Jacala, Hidalgo, 4500 ft., 7 J'J1 1 J vi.25-vii.l.39 (H.H.).
El Banito, Valles, San Luis Potosi, 200 ft., 1 vi. 28.40
(H.H.).

Tancitaro, Michoacan, 6600 ft., 1 J vii.30.40 (H.H.).
Apatzingan, Michoacan, 1050 ft., 1 vii.5.40 (H.H.).

Rio Balsas, Guerrero, 2400 ft., 1 5 v.26.41 (R.P.).
Chilpancingo, Guerrero, 1 $ v.26.41 (R.P.).

The specimens are all in a condition indicating recent emer-
gence.

78. Anteos clorinde Godart.

G. & S., 2 : 148, 726.

R., p. 89, pi. 24g.

Mar., 1944]

Brown: Pierid^:

103

H., p. 656.

Victoria, Tamaulipas, 1 J1 vi.25.35 (H. A. Freeman).

El Pujal, San Luis Potosi, 100 ft., 16 J'J' vii. 17.39
(H.H.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 2 J'J'

iv. 30.41 (R.P.).

Hda. Potrero Viejo, nr. Parare Nuevo, Vera Cruz., 1500
ft., 1 J1 v.5.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 v.10.41 (R.P.).

Jacala, Hidalgo, 4500 ft., 3 $$ vi.2^-vii.7.39 (H.H.).
Apatzingan, Michoacan, 1 5 viii.2.40 (H.H.).

The Apatzingan 5 is badly rubbed, but appears to lack the
usual large yellow area on the forewing.

79. Anteos maerula Fabricius.

G. & S., 2 : 149, 726.

R., p. 89, pi. 24g.

H. , p. 656.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 2 $$ vi.17-18.40 (H.H.).

Victoria, Tamaulipas, 1 vi.25.35 (H. A. Freeman).

60 mi. S. of Victoria, Tamaulipas, 1 J' vii. 6. 36 (H.D.T.).
El Banito, Valles, San Luis Potosi, 200 ft., 4 J'df 2 55
vi. 26.40 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 12 J'J' vii.17.39
(H.H.).

El Sol, Tamazunchale, San Luis Potosi, 150 ft., 1^15

v. 29.41 (R.P.).

Jacala, Hidalgo, 4500 ft., 3 vii.1-10.39 (H.H.).

Chichen Itza, Yucatan, 1 g iv.7.36 (F.M.B.).

The females from El Banito are f. gueneeana Bdv., the one
from El Sol is the nymotypical form.

80. Phoebis sennae Linnaeus.

G. & S., 2 : 141 (as eubule).

R., p. 85.

H. , p. 657.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 3 <?<? 1 5 vi. 16-21.40 (H.H.).

104

Journal New York Entomological Society

[Vol. LIl

Hda. Santa Engracia, Tamaulipas, 2 J'J' vii.27.39
(H.H.).

El Banito, Valles, San Luis Potosi, 400 ft., 2 J'J' 1 5
vi. 26-28.40 (H.H.).

El Sol, Tamazunchale, San Lnis Potosi, 400 ft., 1^1$
v.29.41 (R.P.).

Jacala, Hidalgo, 4500 ft., 3 vi.29.39 (H.H.).

Chichen Itza, Yucatan, 1 § iv.8.36 (P.M.B.) ; 1 §
viii.30.36 (H.D.T.).

Sta. Lucrecia, 1 $ ix. 24.39 (H.D.T.).

Apatzingan, Michoacan, 1200 ft., 2 1 5 viii. 3-5.40

(H.H.).

El Sabino, Uruapan, Michoacan, 1 ^ 1 J vii.15-30.36
(H.D.T.).

Tancitaro, Michoacan, 4500 ft. 1 <£ viii.11.41 (H.H.).

The males from Nuevo Leon are fairly typical of race eubule L.
the others tend toward marcellina Cr., but none of them is as
strongly marked as the typical males from tropical South
America. It is a pity that no one bothers to collect this common
species in sufficient numbers to settle the question of what race
the Mexican material represents. I would designate all of the
above males eubule f. drya Fabricius. I fully realize that else-
where (Amer. Mus. Nov. 368, p. 7, 1929) I stated that Mexican
material was referable to marcellina Cr. I also noted (p. 8) that
the tropical race might well represent two races. The more
material that I see from Mexico the more I feel that these state-
ments need revising, but I have not the material to do so. It is
quite probable that in Mexico the two races so intergrade that we
have a continuous cline from eubule to marcellina.

The El Banito female is f . pallida Cockerell, the El Sol, yamana
Reakirt. The Hda. Vista Hermosa female is a good marcellina
Cr. The Apatzingan female is badly rubbed but is probably
f. yamana. F.M.B.’s from Chichen Itza is pallida while H.D.T.’s
is marcellina.

81. Phoebis philea Linnaeus.

G. & S., 2 : 140, 726.

R., p. 86, pi. 25c.

H. , p. 657.

Mar., 1944]

Brown: Pierim:

105

El Pujal, San Luis P.otosi, 100 ft., 2 vii.17.39 (H.H.).

(H.H.).

Apatzingan, Michoacan, 1200 ft., 1 viii.5.40 (H.H.).

83. Phoebis agarithe maxima Neumoegen.

G. & S., 2: 145, 726 (as agarithe).

R., p. 86 (as argante agaritha).

H. , p .657.

Hda. Vista Hermosa, Nuevo Leon, 1500 ft., 6 3 22

vi.16-21.40 (H.H.).

Sabinas Hidalgo, Nuevo Leon, 960 ft., 2 2

vi. 15-18. 39 (H.H.).

Arroyo del Meco, Tamaulipas, 1320 ft., 2 iv.28.41

(R.P.).

Hda. Santa Engracia, Tamaulipas, 2 J'J' vii.25.39
(H.H.).

Victoria, Tamaulipas, 1 2 vi.28.35 (H. A. Freeman).

60 mi. S. Victoria, Tamaulipas, 1 2 vii.6.36 (H.D.T.).

El Pujal, San Luis Potosi, 100 ft., 3 vii.20-21.39

(H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 £
v.29.41 (R.P.).

Tamazunchale, San Luis Potosi, 500 ft., 1 £ vi.30.40
(H.H.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1
iv.30.41 (R.P.).

f vii.22.39

El Banito, San Luis Potosi, 200 ft., 3 J'J' 1 $ < ^2^40

l (H.H.).

El Mante, San Luis Potosi, 1 iv.29.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 J* v.6.41 (R.P.).

These are all good maxima except the El Sol male which ap-
proaches agarithe. The northern specimens are absolutely typ-
ical, those from the south show some of the brownish markings
on the underside. All the females are f . albarithe Brown.

83c. Phoebis agarithe agarithe Boisduval.

Progresso, Yucatan, 1 2 viii.27.36 (H.D.T.).

Chichen Itza, Yucatan, 1 2 viii.30.36 (H.D.T.).

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Journal New York Entomological Society

[Vol. LII

These two females are much more like typical South American
agarithe than maxima and I have so designated them. They are
not antiilia Brown, which might be expected on the peninsula.
The Progresso specimen is the yellow form, the Chichen Itza one
the white form. Hoffmann has not included this race in his
check-list.

84. Phoebis intermedia Butler.

G. & S., 2: 143, 726 (as rurina).

R., p. 86.

H. , p. 657.

Tancitaro, Michoacan, 4500 ft., 1 vii. 25.40 (H.H.).

86. Aphrissa statira jada Butler.

G. & S., 2 : 147 (as statira).

R., p. 87 (as statira).

H. , p. 657.

El Banito, Valles, San Luis Potosi, 200 ft., 1 J vi.26.40
(H.H.).

This is a pale lemon yellow female like those of true statira.

87. Kricogonia lyside Godart.

G. & S., 2: 150, 151, 726 (as unicolor).

R., p. 89.

H. , p. 658.

Sabinas Hidalgo, Nuevo Leon, 900 ft., 9 10

vi.15-18.39 (H.H.).

Ojo de Agua, Sabinas Hidalgo, N. L., 1000 ft., 2
vi.14.40 (H.H.).

Villa Santiago, Nuevo Leon, 1500 ft., 2 viii.8.39.

(H.H.).

35 km. W. of Linares, Nuevo Leon, 1 J viii.7.39. (H.H.).
Galeana, Nuevo Leon, 2 J'J' 2 22 vii. 28-viii. 4.39. (H.H.).
Hda. Sta. Engracia, Tamaulipas, 1^1} vii.25-27.39
(H.H.).

El Banito, Valles, San Luis Potosi, 1 § vii.22.39 (H.H.).
nr. Apatzingan, Michoacan, 500 ft., 3 J'.J' 3 22 viii.3.40
(H.H.).

Apatzingan, Michoacan, 1050 ft., 1 $ 2 viii.2-5.40

(H.H.).

Mar., 1944]

Brown: Pierid^e

107

El Sabino, Uruapan, Michoacan, 2 1 J yii.15-30.36

(H.D.T.).

Acahuato, Michoacan, 3000 ft., 1 $ viii.2.40 (H.H.).
Tancitaro, Michoacan, 6600 ft., 1 £ vii.20.40 (H.H.).

p 2 cs

It is rather strange that Potts took no specimens of this common
species. The above series contains examples of all of the named
forms that occur in Mexico.

typical $ terrissa $ unicolor $ fantasia

Nuevo Leon 21 5 4 1?

Tamaulipas 1 1

San Luis Potosi 1

Michoacan 7 3 1 4

The specimens noted as fantasia are not typical; they all lack
the costal yellow streak and the ground color of the wings tends
to be greenish rather than yellow. The size of the specimens
varies greatly. This is best shown in the text figure, based upon
the baso-apical radius of the forewings, and the following table
of data.

N. Mean S.D. Critical limits Range

Total

series ... 45 21.37 ± 0.57 mm. 3.66 mm. 14.1 mm., 28.7 mm. 16-28 mm.

Sabinas

Hidalgo 19 22.90 ± 0.34 mm. 3.27 mm. 16.5 mm., 29.4 mm. 16-28 mm.

108

Journal New York Entomological Society

[Vol. LII

88. Eurema albula Cramer.

G. & S., 2: 166.

R., p. 84.

H. p. 658.

K. (4), p. 121.

El Sol, Tamazunchale, San Lnis Potosi, 400 ft., 11
iv.30, v.29.41 (R.P.).

Ojo de Agua, Yera Cruz, 1600 ft., 4 v.12.41 (R.P.).

El Sabino, Urnapan, Michoacan, 1 vii. 15-30. 36

(H.D.T.).

Two of the Ojo de Agua specimens are intermediate to albula
and f. tapeina Bates.

89. Eurema jucunda Boisduval.

G. & S., 2 : 168, pi. 64, f. 9-12.

R., p. 83, pi. 24e.

H. , p. 658.

K., p. 124.

nr. Acapulco, Guerrero, 100 ft., 1 2 v.26.41 (R.P.).
nr. Chilpancingo, Guerrero, 4 jy1 3 2$ v.26.41 (R.P.).
Rio Balsas, Guerrero, 2400 ft., 1 2 v.26.41 (R.P.).

El Sabino, Uruapan, Michoacan, 10 J'J1 9 22 vii.15-30.36
(H.D.T.).

Apatzingan, Michoacan, 3000 ft., 1 $ viii.2.40 (H.H.).
Tancitaro, Michoacan, 6600 ft., 1 y 1 2 vii. 25, viii.11.40
(H.H.).

All the males are of f . sidonia Felder. This is also true of the
females except for two from Chilpancingo which tend toward the
typical in respect to the margin of the hindwings. The general
color of the females varies from white to pale yellow. One of the
Chilpancingo specimens and the one from Rio Balsas have white
forewings and yellow hindwings.

89a. Eurema jucunda lydia Felder.

G. &S., 2:170, pi. 63, f. 20-23.

R., p. 83 (as delia lydia).

H. , p. 658.

K., p. 126 (as palmyra lydia).

Jacala, Hidalgo, 4500 ft., 3 jy 2 2? vi.25-vii.2.39
(H.H.).

Mar., 1944]

Brown: Pierhx®

109

El Pujal, San Lnis Potosi, 100 ft., 5 2 vii.17-

21.39 (H.H.).

Tuxpango, Vera Crnz, 1500 ft., 1 c? 2 5? v-9.41 (R.P.).

Hda. Potrero Viejo, Vera Cruz, 1500 ft., 1 J' 1 J v.5.41
(R.P.).

Orizaba, Vera Cruz, 2000 ft., 4 1 J v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 2 1 2 v.10.41 (R.P.).

This series of “barred” Euremas from the east coast is quite
variable. All have the pearly white under surfaces characteristic
of jucunda. The females are fairly constant, having a reduced
grey bar on the inner margin of the forewing and an incomplete
black margin on the hindwing. One Jacala specimen is faintly
flushed with yellow on the forewing and the Rio Blanco female
bears this flush on the hindwing. The males are extremely varied
but I hesitate to refer the specimen to anything other than lydia.
The marginal band of black on the hindwings is broad and com-
plete on two (Rio Blanco and Orizaba), narrower but complete
on two (El Pujal) and interrupted on the rest. In no case is the
margin reduced to an apical spot. The hindwings range from
yellow (J1 Jacala) to white with many specimens just faintly
tinged with yellow. Until a lot of life history work is done and
enormous series taken over long periods of time for many locali-
ties we will not be able to unravel the puzzle of the “barred”
Euremas.

91. Eurema boisduvaliana Felder.

G. &. S., 2 : 159, pi. 63, f. 1-4.

R., p. 81 (as mexicana, in part).

H. , p. 658.

K, p. 133.

Galeana, Nuevo Leon, 6500 ft., 1 £ 10.29.41 (R.P.).

Jacala, Hidalgo, 4500 ft., 4 1 $?, vi.29-vii.5.39

(H.H.).

El Pujal, San Luis Potosi, 100 ft., 11 vii.12-21.39

(H.H.).

Arroyo del Calabezas, San Luis Potosi, 1 J' iv.30.41
(R.P.).

El Banito, Valles, San Luis Potosi, 1^1$ vi. 26-27. 40
(H.H.).

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Journal New York Entomological Society

[Yol. LII

El Sol, Tamazunchale, San Lnis Potosi, 2 v.29.41

(R.P.).

Fortin, Yera Cruz, 1600 ft., 2 J'J' v.4.41 (R.P.).
Orizaba, Vera Cruz, 2000 ft., 1 v.4.41 (R.P.).

Rio Blanco, Yera Cruz, 2200 ft., 8 J'J' v.10.41 (R.P.).
Chichen Itza, Yucatan, 1 <£ viii.30.36 (H.D.T.).

El Sabino, Uruapan, Michoacan, 1 §?, vii.15-30.36
(H.D.T.).

Apatzingan, Michoacan, 3200 ft., 1 5 viii.2.40 (H.H.).
The questioned female from Jacala is aberrant. The apical
patch on the hindwings is reduced to a few scales along the
nervule. The same is true of the El Sabino female.

92. Eurema xanthochlora Kollar.

G. & S., 2 : 161, 727, pi. 63, f. 5-8.

R., p. 81, pi. 24b.

H. , p. 658.

K., p. 134.

El Banito, Yalles, San Luis Potosi, 200 ft., 1 $ vi.28.40
(H.H.).

Tuxpango, Yera Cruz, 1500 ft., 1 £ v.9.41 (R.P.).

93. Eurema mexicana mexicana Boisduval.

G. & S., 2 : 157.

R., p. 81, pi. 24a.

H. , p. 659.

K, p. 134.

Hda. Yista Hermosa, Villa Santiago, Nuevo Leon, 1500-
3000 ft., 6^1$ vi.17-21.40 (H.H.).

Villa Santiago, Nuevo Leon, 1500 ft., 1 J' viii.8.39
(H.H.).

Jacala, Hidalgo, 4500 ft., 37 J'J' 1 5 vi.23-vii.2.39
(H.H.).

El Pujal, San Luis Potosi, 100 ft., 1 <$ vii.17.39 (H.H.).
El Sol, Tamazunchale, 400 ft., 8 J'.J' v.28-29.41 (R.P.) ;
1$ iv.31.41 (R.P.).

Tuxpango, Yera Cruz, 1500 ft., 1 J v.8.41 (R.P.).

Rio Blanco, Yera Cruz, 2200 ft., 5 v.10.41 (R.P.).

Cumbres (km. 295), Yera Cruz, 6000 ft., 1 v.7.41

(R.P.).

Mar., 1944]

Brown: Pierid^e

111

El Sabino, Uruapan, Michoacan, 3 J'J' vii.15-30.36
(H.D.T.).

Tancitaro, Michoacan, 6600 ft., 1 $ viii.14.40 (H.H.).

Cerro Tancitaro, Michoacan, 7000 ft., 2 vii.20.40

(H.H.).

All these specimens are typical mexicana. Hoffmann includes
in his listing of the species both ab. recta Klots and ab. bieder-
manni Ehrmann with question marks. I have a distinct feeling
that these two “aberrations’7 describe the material from the
northern reaches of the race such as the mountains of Colorado
and should be raised to racial status. Ehrmann’s name is avail-
able for this. There is some variation in size among the speci-
mens before me. A male from Villa Santiago measures only 15
mm. along the baso-apical radius of the forewing while the largest
specimen from El Sol measures 27 mm. The black margin of the
hindwing is variable; on some specimens it is a small patch an-
terior to M2 while in others it extends to Cu2. On these latter
it is broadest at the M2-M3 interspaces and has a basad spur in
Ms— Cul

94. Eurema salome Felder.

G. & S., 2 : 160 (as fabiola).

R., p. 81.

H. , p. 659.

K, p. 136.

Jacala, Hidalgo, 4500 ft., 16 vi.23-vii.2.39 (H.H.).

Fortin, Vera Cruz, 1600 ft., 1 $ v.3.41 (R.P.).

These are all f. limoneus Felder. I disagree with Hoffmann
and agree with Klots on the status of this name. I feel that the
species is splitting into two races and that the northern material
from Mexico is sufficiently different from Ecuadorian salome to
require a name in this genus.

95. Pyristia gundlachia Poey.

G. & S., 2 : 156.

R., p. 81, pi. 21a.

H. , p. 659.

K, p. 137.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft.,2$2vi.l7-22.40 (H.H.).

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Journal New York Entomological Society

[Vol. LII

96. Pyristia proterpia Fabricius.

G. & S., 2: 155.

R., p. 81, pi. 24a.

H. , p. 659.

K., p. 137.

nr. Villagran, Tamaulipas, 1160 ft., 1 J' iv.28.41 (R.P.).
C. Victoria, Tamaulipas, 1 g vi.19.39 (H.H.).

Hda. Sta. Engracia, Tamaulipas, 1 vii.27.39 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 3 1 5 vii. 17-20.39

(H.H.).

Rio Blanco, Vera Cruz, 2200 ft., 2 J'J' v.10.41 (R.P.).
Chichen Itza, Yucatan, 1 g viii.30.36 (H.D.T.).
Acapulco, Guerrero, 100 ft., 1 v.26.41 (R.P.).

Apatzingan, Michoacan, 1050 ft., 5 viii.2-5.40

(H.H.).

El Sabino, Uruapan, Michoacan, 10 J'J' 4 5? vii. 15-30. 36
(H.D.T.).

Tancitaro, Michoacan, 7000 ft., 1^1} vi.25-30.40
(H.H.).

On the northernmost specimen (Villagran) the black markings
are greatly reduced. The Chichen Itza specimen approaches
watsoni from the similar arid area in Ecuador !

97. Pyristia dina westwoodi Boisduval.

G. & S., 2 : 163 (as dina).

R., p. 82, pi. 24b.

H. , p. 659.

K., p. 139.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 7 J'J'
4 22 iv. 30-31, v.28-29.41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1 $
iv.30.41 (R.P.).

Chichen Itza, Yucatan, 1 £ viii.30.36 (H.D.T.) ; 1 2
iv.7.36 (F.M.B.).

N. of Iguala, Guerrero, 1 $ v.22.41 (R.P.).

All the females are much lighter than the males and in many
respects, other than size, compare favorably with f . 2 citrina
Poey from the West Indies. The West Coast male is the exact
counterpart of race parvumbra Kaye from Jamaica!

Mar., 1944]

Brown: Pierid^e

113

98. Pyristia lisa euterpe Menetries.

G. & S., 2 : 162.

R., p. 83.

H. , p. 659.

K., p. 138.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 £ vi.15.39
(H.H.).

nr. Villagran, Tamanlipas, 1150 ft., 3 iv.28.41

(R.P.).

60 mi. So. of Victoria, Tamaulipas, 2 vii.6.36

(H.D.T.).

Jacala, Hidalgo, 4500 ft., 1 vii.6.39 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 1 5 vi.20.39 (H.H.).
Rio Blanco, Vera Cruz, 2200 ft., 1 v.10.41 (R.P.).

The female is f. centralia Herrick-Schaffer {alba Strecker).

99. Pyristia nise perimede Prittwitz.

G. &S., 2:165, 727 (as tenella).

R., p. 83, pi. 24d (as nise).

H. , p. 659.

K, p. 140.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 vi.31.39

(H.H.).

nr. Villagran, Tamaulipas, 1160 ft., 2 J'J' iv.28.41 (R.P.).
C. Victoria, Tamaulipas, 1 J vi.19.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 $ vii.6.36 (H.D.T.).
Hda. Sta. Engracia, Tamaulipas, 1 J' vii.25.39 (H.H.).
Jacala, Hidalgo, 4500 ft., 5 2 5? vi.23-vii.2.39

(H.H.).

El Pujal, San Luis Potosi, 100 ft., 3 vii.18-21.39

(H.H.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1 J*

iv. 30.41 (R.P.). z

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1^1$

v. 28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 150 ft., 5 2 5?

vi. 26.40, vii.22.39 (H.H.) ; iv.30.41 (R.P.).

Hda. Potrero Viejo, Paraje Nuevo, Vera Cruz, 1500 ft.,
3 v.5.41 (R.P.).

114

Journal New York Entomological Society

[Vol. LII

Orizaba, Vera Cruz, 2000 ft., 2 v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 8 v.10.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 5 v.12.41 (R.P.).
Acapulco, Guerrero, 100 ft., 1 J' v.23-25.41 (R.P.).

Sta. Lucrecia, 1 ix.24.39 (H.D.T.).

Apatzingan, Michoacan, 500 ft., 1 <$ viii.3.40 (H.H.).

El Sabino, Uruapan, Michoacan, 5 2 vii.15-30.36

(H.D.T.).

There is a great deal of variation in this series, none of it re-
lated to geographic distribution. The ground color varies from
pale to deep lemon. The dark margin on the hindwings varies
from complete to absent ; on over half of the specimens it is rep-
resented by only a row of black dots on the nervules. There is
some variation in the extent of the black marking on the fore-
wing. On some the underside is boldly marked; on others it is
almost immaculate.

100. Abaeis nicippe Cramer.

G. & S, 2 : 155, 727.

R., p. 81, pi. 24a.

K., p. 132.

H. , p. 659.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 5 J'J' vi.15-16.39
(H.H.).

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1300 ft.,
1 vi.14.40 (H.H.).

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 1 <$ 2 ?? vi.16-21.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 2 jy1 vii.30-viii.3.39
(H.H.).

Jacala, Hidalgo, 4500 ft., 11 J'J' vi.23-vii.2.39 (H.H.).
El Pujal, San Luis Potosi, 100 ft., 1 J1 n.d. (H.H.).
Hda. Potrero Viejo, nr. Paraje Nuevo, Vera Cruz, 1500
ft., 1 $ vi.5.41 (R.P.).

Monte Alban, Oaxaca, 6000 ft., 1 y v.18.41 (R.P.).

The Monte Alban specimen is notable for the reduction of the
dark apical markings on it.

101. Nathalis iole Boisduval.

G. & S., 2 : 172, pi. 64, ff. 15-18.

Mar., 1944]

Brown: Pierid^

115

R., p. 95, pi. 27h (as jole).

H., p. 659.

bet. Adjuntas & Sabinas Hidalgo, Nuevo Leon, 3000
ft., 1 c? vi.21.40 (H.H.).

Sabinas Hidalgo, Nuevo Leon, 1500 ft., 1 1 J vi.17.40

(H.H.).

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1000 ft.,
1 g vi. 15.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 vii. 30.39 (H.H.).

nr. Villagran, Tamaulipas, 1160 ft., 1 iv.28.41 (R.P.).

C. Victoria, Tamaulipas, 1 § vi.19.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 2 vii.6.36
(H.D.T.).

Jacala, Hidalgo, 4500 ft., 10 1 5 vi.23-vii.l.39.

(H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 $
v.29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 150 ft., 2 ££ iv.30.41
(R.P.).

El Pujal, San Luis Potosi, 100 ft., 1^1} vii.l7.39J
(H.H.).

Tuxpango, Vera Cruz, 1500 ft., 2 ££ v.9.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 g 1 2 v.6.41 (R.P.).

Cumbres (km. 295), Vera Cruz, 8000 ft., 3 ££ v.7.41
(R.P.).

Monte Alban, Oaxaca, 6000 ft., 1 2 v.18.41 (R.P.).

The females from all localities other than Jacala and Monte
Alban are f. irene Pitch. One of the Jalaca males has the fore-
wing bar reduced to a submarginal spot.

(Pierini)

103. Neophasia terlootii Behr.

R., p. 55, pi. 18a.

H., p. 660.

Cerro Tancitaro, Michoacan, 5500 ft., 1 $ viii.17.40
(H.H.).

This capture of Hoogstraal’s extends the range of this species-
over 300 miles southward. I suspect that it will be recovered still

116

Journal New York Entomological Society

[Vol. LII

further south, possibly in Central America in the pine-oak forests
of Guatemala and Honduras.

104. Catasticta flisa Herrick-Schaffer.

G. & S., 2 : 117.

R., p. 73, pi. 22e.

H. , p. 660.

Jaeala, Hidalgo, 4500 ft., 1 vi.29.39 (H.H.).

This extends the range of the species northward from Vera
Cruz (see H., p. 660).

105. Catasticta nimbice nimbice Boisduval.

G. & S., 2 : 118.

R., p. 70, pi. 22b.

H. , p. 660.

Jaeala, Hidalgo, 4500 ft., 2 vi.2L-28.39 (H.H.).

Tancitaro, Michoacan, 6600 ft., 1 J vii.30.40 (H.H.).
The female is freshly emerged, the males are worn.

110. Appius ilaire Godart.

G. & S., 2 : 135 (as margarita).

R., p. 59, pi. 19c.

H. , p. 661.

El Banito, Valles, San Luis Potosi, 200 ft., 1 £ vi. 28.40
(H.H.).

113. Pieris protodice Boisduval & Le Conte.

G. & S., 2 : 130, 724.

R., p. 59.

H. , p. 661.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 5 vi.15.39
(H.H.).

Hda. Vista Hermosa, nr. Villa Santiago, Nuevo Leon,
1600 ft., 1 3 vi.16.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 J viii.1.39 (H.H.).
Monterrey, Nuevo Leon, 1600 ft., 1 § iv.27.41 (R.P.).
nr. Villagran, Tamaulipas, 1160 ft., 1^1} iv.28.41
(R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 J'J' vi.28.40
(H.H.).

Mar., 1944]

Brown: Pierid^e

117

114. Pieris rapae Linnaeus.

R., p. 58, pi. 19b.

H., p. 661.

Galeana, Nuevo Leon, 6500 ft., 2 <£<§ viii.4.39 (H.H.).
This species seems to have reached Mexico since the publication
of the Biologia (since 1900?).

116. Leptophobia aripa Boisduval.

G. & S., 2 : 136, 726 (as elodia).

R., p. 60, pi. 19d.

H. , p. 661.

nr. Villagran, Tamaulipas, 1160 ft., 1 5 iv.28.41 (R.P.).
Jacala, Hidalgo, 4500 ft., 2 J'J' vi.28.39 (H.H.).

Fortin, Vera Cruz, 1600 ft., 1 J1 v.3.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 v.12.41 (R.P.).

Orizaba, Vera Cruz, 200 ft., 1 £ v.6.41 (R.P.).

117. Itaballia demophile calydonia Boisduval.

G. & S., 2 : 131.

R., p. 63.

H. , p. 661.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 £
v.29.41 (R.P.).

The subapical cross bar on the upper sides of the forewings is
totally lacking, on the underside it is reduced to little more than
a broad line. This specimen extends the range northward from
southern Vera Cruz (see H., p. 661).

119. Pieriballia viardi viardi Boisduval.

G. & S., 2 : 133, 725, pi. 62, f. 1-4.

R., p. 62.

H. , p. 661.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 $
iv.31.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 J* 1 J vii.20-21.39
(H.H.).

121. Ascia monuste cleomes Boisduval.

G. & S., 2 : 132, 725 (as monuste).

R., p. 57.

H. , p. 662.

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Journal New York Entomological Society

[Vol. LII

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1600
ft., 1 ? vi.17.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 5 iv.29.41 (R.P.).

nr. Villagran, Tamaulipas, 1160 ft., 1 § iv.28.41 (R.P.).

Arroyo del Meco, Tamaulipas, 1 y iv.28.41 (R.P.).

C. Victoria, Tamaulipas, 2 §f vi.27.35 (H. A. Free-
man) ; 2 yy vi.19.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 3 yy vii.6.36
(H.D.T.).

Jacala, Hidalgo, 5400 ft., 1 y 1 ? vii.1.39 (H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 yy

v. 29.41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1 y
10.30.41 (R.P.).

El Baiiito, Valles, San Luis Potosi, 200 ft., 2 yy 2

vi. 26.40, vii.19.39 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 3 yy vii. 17-19. 39
(H.H.).

Tuxpango, Vera Cruz, 1500 ft., 1 y v.9.41 (R.P.).

Hda. Potrero Viejo, Paraje Nuevo, Vera Cruz, 1500 ft.,
1 ? v.5.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 y v.10.41 (R.P.).

Apatzingan, Michoacan, 1200 ft., 2 yy 1 J viii.5.40
(H.H.).

The males are rather uniform throughout. The marginal dark
spots on the hindwing are variable, being represented by either
a few scales or a small triangular patch. This does not seem to
be related to the origin of the specimens. One male from 60
miles South of Victoria is almost immaculate on both upper and
lower surfaces. The females fall into two clearly defined groups.
Those from Nuevo Leon, Tamaulipas and Michoacan have a pink-
ish iridescence on the upper surface and one (from Victoria)
approaches phileta Fabricius. Those from San Luis Potosi,
Hidalgo and Vera Cruz are a dull buff -yellow on the hindwings
and the same with a slightly pinkish tone on the forewings.

123. Melete isandra Boisduval.

G. & S., 2 : 139, 726, pi. 61, ff. 13, 14.

R, p. 77, pi. 23b.

Mar., 1944]

Brown: Pierid^

119

H., p. 662.

El Banito, Valles, San Lnis Potosi, 200 ft., 1 $ 1 J
vii.19.39 (H.H.).

El Pnjal, San Lnis Potosi, 100 ft., 3 yii. 17-20.39
(H.H.).

BIBLIOGRAPHY

(1) . Godman & Salvin, Biologia Centrali-Americana, Rhopalocera, 2: 113-

188. 1889-1890.

(2) . Roeber, in Seitz’ Macrolepidoptera of the World, vol. 5, pp. 53-111.

1909.

(3) . Hoffmann, Anales del Instituto de Biologia, 11: 639-739. 1940.

(4) . Klots, Entomologica Americana, 9: 99-171. 1929.

No. 2

Vol. LII

JUNE, 1944

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B. WEISS

Publication Committee

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.

LANCASTER, PA.

NEW YORK, N. Y.

1944

CONTENTS

A Generic Review of the Subfamily Phyllobaeinae

By A. B. Wolcott 121

The Reproductive Functions of the Army-Ant Queen as
Pace-Makers of the Group Behavior Pattern

By T. C. Schneirla 153

A Re-Survey of Papaipema Sm. (Lepidoptera)

By Henry Bird 193

Insect Introductions and War

By William M. Boyd 200

Catalogue of North American Psychodidse

By William F. Rapp, Jr 201

Recent Work by the International Commission on Zoologi-
cal Nomenclature

By Francis Hemming 211

NOTICE: Volume LII, Number 1, of the Journal of the
New York Entomological Society was published on
April 6, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, I’a..
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LII June, 1944 No. 2

A GENERIC REVIEW OF THE SUBFAMILY PHYLLO-
BiENINiE (OLIM HYDNOCERIN^E) (COL.)

By A. B. Wolcott
Downers Grove, III.

It has become increasingly evident that the genera of this sub-
family are in a very chaotic condition, and that they are neither
clearly defined nor readily understood. This state of confused
conditions has been produced by various causes, such as the inclu-
sion within the group of extraneous elements, which should have
been assigned elsewhere ; the omission of certain genera ; disre-
gard of the rules of priority and nomenclature; the all too fre-
quent totally inadequate diagnoses and the lack of an ample and
accurate synonymy, all of which have been contributing factors
tending to create the utmost disorder.

In the present paper the writer will endeavor to eliminate the
alien elements, to tabulate the several genera which are compo-
nents of this subfamily, to give where needful, or at all possible
an amended diagnosis, to give an ample synonymy, to indicate
the type of each genus and record their geographic range.

Phyllob^enin.® Subfamily nom. nov.

Subfamily Characters

Cleridae; maxillary palpi with terminal segment cylindrical,
usually much shorter than that of the labial palpi, which are
large, dilated and securiform. Head with the eyes included
broader than the prothorax; antennae short; eleven-segmented
(with the exception of one or two genera), usually with a two-

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[Vol. LIT

to five-segmented clava (rarely feebly developed). Thorax with
the sides more or less dilated at, or before the middle. Elytra
usually elongate, completely covering the abdomen, or more or
less abbreviated, elytral sculpture variable. Terminal segments
of abdomen, especially in the male, with sexual modifications.
Legs long, slender, tarsi five-segmented, basal segment very small,
atrophied or wanting, when present covered by the second seg-
ment, or elongate and distinct, not at all hidden from view from
above.

The following key to the genera, studied in connection with the
generic diagnosis which follow, will, it is believed, render possible
a ready recognition of the several genera.

KEY TO THE GENERA

10.

11.

Antennae ten-segmented, clava uni-segmented Lasiocallimerus Corp.

Antennae eleven-segmented 2.

Antennal clava indistinct 18.

Antennal clava distinct; basal segment of hind tarsi small, atrophied or

absent, usually covered by the second segment 3.

Clava two-segmented 4.

Clava three- to five-segmented ......: 8.

Terminal segment of clava larger than the tenth Parmius Sharp.

Terminal segment of clava subequal in size to tenth, sometimes connate,

causing the antennae to appear ten-segmented Neohydnus Gorh.

Terminal segment of clava minute or very small 5.

Tarsal ungues with a broad basal tooth Phyllobcenus Dep**

Tarsal ungues simple or at most only slightly thickened at base 6.

Prothorax much longer than broad; third segment of antennae about

twice as long as broad, cylindrical ! Isohydnocera Chpn.

Prothorax subequilateral , , 7.

Third segment of antennae stout, only slightly longer than broad.

Emmepus Motsch.

Third segment of antennae as broad or broader than long, trapezoidal.

Wolcottia Chpn.

Clava three-segmented; basal segment of hind tarsi short 9.

Clava three- to five-segmented; basal segment of hind tarsi long, not

covered by the second segment 15.

Clava rather indistinct; protliorax one-half longer than broad.

Cephaloclerus Kuw.

Clava distinct .....: - 10.

Body apterous; eyes coarsely granulate Paupris Sharp.

Body alate; eyes finely granulate 11.

Eyes flat or depressed Abrosius Fairm.

Eyes strongly convex : 12.

June, 1944]

Wolcott: Phyllob^nin^e

123

12.

13.

14.

15.

16.

17.

18.

Eyes large, globular, entire Isolemidia Gorh.

Eyes smaller 13.

Tarsi of hind legs nonlamellate Metaxina Broun.

Tarsi of hind legs with at least two segments lamellate 14.

Clava lax, serrate I Eurycranium Blanch.

Clava rather compact; eyes feebly emarginate Lemidia Spin.

Very small (3 mm.), Phyllobcenus- like in form Theano Cast.

Much larger (5 mm. or more) : 16.

Form broad and compact; elytra never with scales; clava five-seg-
mented sub. gen. Brachycallimerus Chpn.

Form elongate or oblong; adorned with white or yellow scales 17.

Head, the eyes included, only slightly broader than the anterior part of
the prothorax sub. gen. Callimerus Gorh.

Head, the eyes included, much broader than the anterior part of the
prothorax; body depressed sub. gen. Cucujocallimerus Pic.

Elytra depressed, much broader than the prothorax at base, elongate.

Evenus Cast.

Elytra convex, elongate, only slightly broader than the prothorax at
base Allelidea Waterh.

Genus Phyllob^inus Dejean

A hundred and twenty years ago Thomas Say (Jonr. Acad.
Phila., Ill, 1823, p. 192), described an insect to which he gave
the name Clerus humeralis, a year later E. F. Germar (Ins. Spec.
Nov., I, 1824, p. 80) described the same insect (which had evi-
dently come into his possession bearing a label with the name
given to the species by Say), as he adopted for his species the
name of Clerus humeralis , thus creating a synonym of Say’s
species. Several years later Count Dejean (Cat. des Col. de la
coll., edition 3, 1837, p. 127) included under Phyllol>cenus Dejean,
humeralis Germ., which is the first and only valid species, con-
tained in the list of ten names, nine of which are given with
“Dej.” as author, but as he did not describe any of these species
they are to be considered as nomina nuda. Dejean’s citation of
humeralis Germ., according to the International Code ( vide
Article 21 and Opinion 1, B) constitutes an indication, and gives
validity to the genus Phyllobcenus Dejean, with P. ( Clerus )
humeralis Say as the genotype.

Edward Newman (1838: 379) erected the genus Hydnocera
based on serrata Newm., which is a synonym of pallipennis Say
(1825: 176). Newman two years later (1840: 362) described

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three more species of this genus, at which time he states that
“The Count Dejean, in his ‘Catalogue des Coleop teres’ has given
to the genus the provisional name of Phyllobaenus : this I learn
by finding one of them so named by Dr. Harris.” He also re-
marks ‘ ‘ One of the species has been described by Germar, in his
‘Insectorum Species,’ under the name of Clems humeralis ; and
the same insect has also been labeled as the Tillus humeralis of
Say, but I have no reference whatever to any description by
the American entomologist.” Newman’s statements show how
deeply he was imbued with the antiquated and obsolete idea that
a generic name unaccompanied by a characterization or diag-
nosis was of no significance. And upon this supposition some
of these names have been freely and rashly employed as names
to designate other genera. An instance of the kind just men-
tioned is that of Phyllohcenus Spinola (Clerites II, 1844, p. 1),
of which he makes the following statements: “My Phyllobenes
are not those of Mr. Dejean, the latter belong to the G. [enus]
Hydnocera of Mr. Newman, a genus of which that author has
published the characters and of which he is the true founder,
the name that he has assigned to it has incontestably the right of
priority. That of Phyllohcenus having become vacant, I like it
better to take possession of it than rack one ’s brains in looking for
a new name to apply to a species that Mr. Dejean has placed in
the G. [enus] Notoxus and which cannot remain there.” The
action of the present writer in assigning Phyllobcenus to its
proper nomenclatural status leaves Phyllohcenus Spinola vacant,
and necessitates a new name in its stead, the name Phlogisto-
sternus (/Aoyiotos, crematus areppov, sternum, is here proposed.

Genotype. — Phyllobcenus dislofatus.

The writer’s thanks are due his good friend Dr. Edward A.
Chapin, Curator Division of Insects, United States National Mu-
seum, who has assisted the author in many ways; also it was he
who first called the author’s attention to the fact that Hydnocera
Newman is a synonym of Phyllobcenus Dejean.

Phyllohcenus Dejean, Cat. des Col., 3rd Edition, 1837, p. 127.

Hydnocera Newman, The Ent. Mag., Lond., V, 1837 (1838), p.
379. — Newman, Mag. Nat. Hist., (2) IV, 1840, p. 362. — Spinola,
Rev. Zool., 1841, p. 75. — Klug, Clerii, 1842, p. 311. — Spinola, Mon.

June, 1944]

Wolcott: Phyllob^enin^e

125

Clerites, II, 1844, p. 39. — Leconte, Ann. Lyc. Nat. Hist. New
York, V, 1849, p. 26. — Lacordaire, Gen. Col., IV, 1857, p. 471. —
Chenu, Encycl. d’Hist. Nat. Col., II, 1860, p. 268. — Leconte,
Classif. Col. N. Amer., I, 1861, p. 196. — Gorham, Trans. Ent. Soc.
Lond., 1877, pt. 3, p. 260. — Gorham, Biol. Centr.-Amer., Ill, pt. 2,
1883, p. 168. — Leconte and Horn, Classif. Col. N. Amer., ed. II,
1883, p. 218. — Wickham, Can. Ent., XXVII, 1895, pp. 245 and
251. — Schenkling, Gen. Ins. (Wytsman) Cler., 1903, p. 92. —
Schenkling, Col. Cat. (Jnnk) Cler., pt. 23, 1910, p. 102. — Gahan,
Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. — Wolcott, Bull. Ind.
Dept. Geol. Nat. Res., I, 1910, p. 855. — Chapin, Bull. Brookl. Ent.
Soc., XII, 1917, p. 83.— Chapin, Proc. Biol. Soc. Wash., XXXV,
1922, p. 55. — Bradley, Man. Gen. Beetles Amer. N. of Mex., 1930,
p. 105. — Chagnon, Le Nat. Canad., LXII, 1935, pp. 174 and 175.

Theano Chevrolat, Ann. Soc. Ent. France, I (2), 1843, p. 33.

Diagnosis of Phyllobcenus Dejean

Head, with the eyes included, broader than the prothorax,
mandibles feebly curved with a strong internal tooth before the
apex. Head very short, front large, vertical, eyes rather large,
prominent, oval, transverse, entire or the front margin with a
feeble vestige of emargination. Antennas very short, eleven-seg-
mented, first segment thick, obconical, second very short, seg-
ments three to nine nearly cylindrical, the ninth segment as
broad as long, tenth segment large, swollen, ovate, terminated
by the scarcely discernible, minute eleventh segment. Maxillary
palpi with terminal segment short, nearly cylindrical or atten-
uate, apex truncate, terminal segment of labial palpi large, trans-
versely securiform. Prothorax usually longer than broad, sides
more or less dilated before the middle. Elytra much broader
than the prothorax at base, usually moderately elongate parallel
or narrowed near the middle, often abbreviated at apex, sculp-
ture variable. Legs long, posterior femora reaching slightly be-
yond the apex of the elytra, tarsi five-segmented, basal segment
short, the three succeeding lamellate, the posterior tarsi with the
lamella of basal segment long and compressed, lamellae of second
and third segments depressed, bifid. Ungues with a broad basal
tooth.

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[Yol. LII

Newman in his diagnosis of the genus Hydnocera states that
the antennae are 10-articulate, but it is evident that he failed to
observe the very small and closely united true apical segment.
This error was corrected by Lacordaire (1857: 471).

Genotype. — Phyllobcenus ( Clerus ) humeralis Say. Monobasic.

Geographical range, 118 species of North, Central and South
America.

Genus Lemidia Spinola

Lemidia Spinola, Rev. Zool., 1841, p. 75. — Klug, Clerii., 1842,
p. 311. — Spinola, Clerites, II, 1844, p. 32. — Lacordaire, Gen. Col.,
IY, 1857, p. 470.— Chenu, Encycl. d’Hist. Nat. Col., II, 1860, p.
267. — Gorham, Trans. Soc. Ent. Lond., 1877, p. 249. — Schenkling,
Deutsch. Ent. Zeit., 1898, p. 169. — Schenkling, Gen. Ins., (Wyts-
man) Cler., 1903, p. 90. — Schenkling, Col. Cat., (Junk) Cler.,
1910, p. 108. — Lea, Ann. Soc. Ent. Belg., LI, 1907, pp. 331 et 362.

Eumede Pascoe, Ann. Mag. Nat. Hist., (4) XVII, 1876, p. 50. —
Broun, Man. New Zealand Col., I, 1880, p. 334.

Hydnocera Newman, The Entom., 1841, p. 36. — Newman, idem,
1842, p. 365. — White, Cat. Cleridae Brit. Mus., IV, 1849, pp. 61
et 62.

Clerus Newman, Zoologist, 1843, p. 119 (sp. hilaris).

Hoploclerus White, l.c. p. 40. — Westwood, Proc. Zool. Soc.
Lond., XX, 1852, p. 52.

Hydnocerus Westwood, lac cit., 1852, p. 47.

Lemidius Westwood, idem, 1852, p. 47.

Laiomorphus Pic, Mel. Exot.-ent., L, 1937, p. 20.

Diagnosis of Lemidia, revised

Body small, rather elongate, parallel or posteriorly wider.
Head short, with the eyes broader than the prothorax, front
broad between the eyes, which are large, prominent, feebly, shal-
lowly emarginate in front, finely granulated. Maxillary palpi
with the terminal segment cylindrical, somewhat oval, acute at
apex ; that of the labial palpi much larger, depressed, securiform.
Labrum prominent, emarginate. Antennae very short, reaching
the hind margin of the head, eleven-segmented, segment 1 thick
and long, subcylindrical, not rising to the height of the front, seg-
ment 2 very short, swollen at middle and subglobular, segments

June, 1944]

Wolcott: Phyllob^enin^e

127

3 to 8 short, diminishing in length, without becoming sensibly
narrower, segments 9 to 11 thick and depressed, equally dilated,
forming a quite distinct clava of these three segments, the first two
are equal, subtransverse, the last longest and oval. Prothorax
longer than broad, narrower than the head, much narrower at
base, anteriorly with at most a feeble transverse impressed line.
Elytra oblong, surface depressed, entirely covering the abdomen,
irregularly punctured, suture usually closed, spices conjointly
rounded, rarely separately mucronate. Legs long and slender,
the femora extending to or beyond apex of the elytra, tarsi five-
segmented (four segments only visible from above), the first
segment very small or rudimentary but usually perfectly visible
from below, the first three lamellate, the second, third and fourth
segments diminishing in length. Ungues small, simple or at
most with an indistinct tooth at base.

Genotype. — Lemidia ( Hydnocera ) nitens Newman (Spinola).
Monobasic.

Geographical range, sixty-two species of this genus are found
in Australia, one in New Zealand, two in Tonkin, and seven in
Tasmania.

Genus Isohydnocera Chapin

Isohydnocera Chapin, Bull. Brookl. Ent. Soc., XII, 1917, p.
83. — Bradley, Man. Gen. Beetles Amer. N. of Mex., 1930, p. 105.

Diagnosis of Isohydnocera

“Body very elongate; front vertical; eyes prominent, finety
granulate, entire or minutely emarginate near antennal inser-
tion; terminal segment of maxillary palpi cylindro-conical, that
of labial palpi large, triangular; mandibles with a tooth near
apex on the inside. Antennae 11-segmented, first segment thick,
moderately long and somewhat arcuate, second short, globose,
third to ninth cylindrical, longer than broad, tenth very large,
forming with the eleventh a compact club which is much thicker
than the ninth segment. Prothorax distinctly longer than broad,
nearly cylindrical, lateral dilation weak. Elytra broader than
the thorax, shorter than the abdomen, attenuate or parallel and
truncate, lateral margins and usually tips strongly serrate. Legs
long and thin, tarsi with five segments, the first covered by the

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Journal New York Entomological Society

[Vol. LII

second from above, segments two, three and four bearing lamel-
lae beneath, ungues long and thin, simple” (Chapin).

■ ‘ To this genus are to be assigned the following species : I. curti-
pennis Newman ( longicollis Ziegl.), tabida Lee., schusteri Lee.,
ornata Wole., gerhardi Wole., pusilla Schaeff., aegra Newm.,
brunnea sp. nov. and albocincta Horn” (Chapin).

Genotype. — Isohydnocera ( Hydnocera ) curtipennis Newman,
designated by Chapin.

Geographical range, North, Central and South America.

Genus Wolcottia Chapin

Wolcottia Chapin, Bull. Brookl. Ent. Soc., XII, 1917, p. 84.—
Bradley, Man. Gen. Beetles Amer. N. of Mex., 1930, p. 105.

Diagnosis of Wolcottia

“Body elongate, slightly attenuate posteriorly. Head short,
front flat, lateral margins slightly concave to accommodate eyes,
which are not prominent, finely granulate and minutely cemargi-
nate as in Isohydnocera. Labrum broad, entire. Mandibles
falciform, with a small internal tooth near apex. Terminal seg-
ment of maxillary palpi conical, as long, or slightly longer than
the preceding, that of the labial palpi very large, triangular.
Antennae short and stout, 11-segmented ; first segment slightly
longer than broad, second nearly spherical, third to eighth as
long as broad, trapezoidal, becoming progressively broader, ninth
broader than long, nearly as wide as tenth, tenth and eleventh
forming an oval mass nearly as long as the three preceding seg-
ments. Thorax slightly broader than long, sides feebly dilated,
slightly narrower than the head with eyes. Elytra at humeri
broader than the head across eyes, attenuate posteriorly, tips
rounded separately and with irregular margins. Legs long and
thin, hind femora reaching beyond the tips of the elytra, tarsi
of five segments, the first covered by the second from above,
second, third and fourth bearing lamellae beneath. Ungues long
and thin, simple or but slightly thickened” (Chapin).

Genotype. — W. ( Hydnocera ) pedalis Leconte, designated by
Chapin.

Geographical range, North America.

June, 1944]

Wolcott: Phyllob^enin^e

129

Dr. Chapin states that “The three species assigned to this
genus, W. pedalis Lee., sobrina Fall, and parviceps Schaeffer,
have antennae which seem to be midway between the three-seg-
mented club type of Lemidia Spinola and the two-segmented club
type of Hydnocera Newman.”

Genus Parmius Sharp

Parmius Sharp, Ent. Mo. Mag., XIII, 1877, p. 272. — Broun,
Man. New Zealand Col., I, 1880, p. 331. — Schenkling, Gen. Ins.,
(Wytsman) Cler., 1903, p. 96. — Schenkling, Col. Cat., (Junk)
Cleridae, pars 23, 1910, p. 112.

Diagnosis of Parmius, revised

Elongate, alate, slender. Head with the eyes as broad as the
elytra, broader than the thorax. Terminal segment of the maxil-
lary palpi small, that of the labial palpi very large and broad.
Eyes very prominent, finely granulated, entire or with a more
or less definite emargination, antennae short and rather slender,
11-segmented, the two or three terminal segments forming a short,
broad clava, the tenth being strongly transverse and the eleventh
rather stouter than the tenth and hardly as long as broad.
Thorax subcylindrical but a little dilated at the middle, about as
long as broad, but narrower than the elytra, constricted in front
and behind, very narrow and parallel, the humeri distinct and
rectangular. Legs long, tarsi moderately long and broad, very
distinctly five-segmented, the second segment inserted very near
the tibia. Hind coxae rather widely distant.

“The two species ( debilis Shp., and longipes Shp.,) differ from
Paupris aptera by their finely granulated eyes, by the presence
of wings and the more normal form of the elytra ’ ’ ( Sharp ) .

Genotype. — Parmius longipes Sharp, by present designation.

Geographical range, four described species known, all from
New Zealand.

Genus Neohydnus Gorham

Neohydnus Gorham, Ann. Mus. Genova, (2) XII, 1892, p.
742. — Schenkling, Gen. Ins., (Wytsman) Cler., fasc. 13, 1903,
p. 94. — Schenkling, Col. Cat., (Junk) Cleridae, pars 23, 1910,
p. 107.— Chapin, Phil. Jour. Sci., XXV, No. 2, 1924, p. 180.

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Journal New York Entomological Society

[Yol. LIJ

Emmepus Motchoulsky, Bull. Imp. Nat. Moscou, XXXIV,
1861, p. 127 (sp. flavipes).

Diagnosis of Neohydnus, revised

Head vertical. Labrnm entire ; mandibles stout, falcate, with
a short internal tooth toward apex. Eyes large, finely granu-
lated, pubescent, entire or minutely emarginate at base of an-
tennae; the head plane between the eyes (very much as in Cal-
limerus). Terminal segment of maxillary palpi cylindro-acumi-
nate, that of labial palpi dilated, elongate-triangular. Antennae
short 11-segmented (often apparently ten-segmented, the tenth
and eleventh segments being anchylosed), first segment short
and thick, slightly bent, second nearly spherical, third to ninth
trapezoidal, becoming successively shorter to eighth or ninth,
ninth sometimes longer than the eighth, tenth and eleventh to-
gether oval and flattened. Thorax with length and breadth sub-
equal, not widened in front (as in Phyllobcenus) , with a con-
stricted line near the front, sides slightly dilated below this con-
struction, lateral foveae deep, oblique above dilations, usually
two or three in number, no basal constriction but margined at
base, anterior coxal cavities widely open behind. Elytra usually
as long as body, confusedly punctate, no trace of striae, suture
slightly dehiscent toward apices, lateral margin and tips usually
strongly serrate. Abdomen with six visible ventral segments.
Sexual modifications of males usually conspicuous. Legs slender,
femora somewhat swollen ; tarsi short, of five segments, first seg-
ment short and ventral to second, ungues with a broad basal
process.

Genotype. — Neohydnus despectus Gorham.

Geographical range, Indo-Malaysia.

Genus Metaxina Broun

Metaxina Broun, Ann. Mag. Nat. Hist., (8) III, 1909, p. 407.

Diagnosis of Metaxina, revised

Head (eyes included) nearly as large as the thorax, forehead
limited between the antennae by a slightly raised suture. Palpi
stout, long, the terminal segment of maxillary palpi quite oblique

June, 1944]

Wolcott: Phyllob^nin^e

131

at apices; that of the labial palpi subtriangular or securiform,
truncate at apices. Eyes prominent, distinctly granulated, ap-
parently rotundate, in reality transversely broadly oviform, en-
tire. Antennae eleven-segmented, inserted not in cavities, but
on slight prominences, or articulations in front of the eyes ; basal
segment stout, pyriform, third segment slightly longer than
second or fourth ; segments four to eight nearly equal in breadth,
each longer than broad, and narrowed at the base ; segments nine
and ten distinctly broader than the preceding, but little longer ;
eleventh rather larger and oviform ;. these organs therefore more
nearly resemble those of Paupris and Parmius than of Phymato-
phcea. Prothorax somewhat cordiform, transverse, apex trun-
cate, base rounded, the lateral margins fine and curved toward
the base, but without forming posterior angles; scutellum elon-
gate. Prosternum truncate in front. Coxae with trochanters;
the anterior prominent, separated only by the prosternal process ;
intermediate rather less prominent, almost contiguous; the pos-
terior small, only moderately separated. Metasternum moder-
ately elongate, its sides distinctly punctured. Elytra parallel-
sided, broader than the thorax, with obtusely prominent humeri,
the punctuation moderately coarse, not close and subseriate, sur-
face subdepressed, apices not quite smooth, elongate. Tarsi five-
segmented, basal segment abbreviated above, its lobes prolonged
below; segments two to four of nearly equal length, their lobes
also prolonged but without perceptible lamellae underneath ; fifth
simple, nearly as long as the preceding four together; basal seg-
ments of the anterior tarsi furnished with long slender, brush-
like setae below. Abdomen composed of six segments, the basal
segment rather larger than the others, segments two to five about
equal ; sixth short, deeply emarginate, with a supplementary
conical segment protruding therefrom.

Genotype. — Metaxina ornata Broun, designated by Broun.

Geographical range, 1 species of New Zealand.

Broun remarks that “the type of this genus is quite unlike
any Australian or New Zealand species known to me, and cer-
tainly an aberrant form, owing to the absence of tarsal lamellae,
the unnotched eyes, &c.” This genus and its genotype has not
been mentioned in the literature since the genus was created by

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Broun. This is truly an anomalous genus, but this being an
insular insect, and obviously a highly adaptive form, the singu-
larity of its aspect need cause no great doubt as to the propriety
of including this genus in the present subfamily.

Genus Abrosius Fairmaire

Abrosius Fairmaire, Bull. Mus. Hist. Nat., Paris, VIII, 1902,
p. 315. — Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 96. —
Schenkling, Col. Cat., (Junk) Cler., 1910, p. 112. — Gahan, Ann.
Mag. Nat. Hist., (8) V, 1910, p. 65.

Diagnosis of Abrosius translated from the original

“This new genus of Cleridae approaches the Hydnocerides by
the tarsi having the first segment indistinct, the eyes entire and
the antennae inserted near their lower border; but the eyes are
rather depressed, the legs of ordinary length and the facies is
very different from that of Evenus, recalling greatly that of
Aulicus. The antennae are slender, of eleven articles, the three
terminal segments forming a small, rather narrow, lax mass ; the
front is rather large, but the eyes moderately convex. Prothorax
scarcely longer than broad, base moderately narrow, sides
rounded, scarcely sinuate at base. Elytra oblong, humeri rather
angulate, surface with basal portion rather strongly seriate-
punctate, posteriorly minutely and confusedly punctate. Legs
slender, rather short, tarsi seen from above four-segmented, the
first segment indistinct, the intermediate segments provided with
rather long lamellae. ’ ’

Genotype. — Abrosius cyaneorufus Fairm., by sole inclusion.

Geographical range, 1 species of Madagascar.

The genus Abrosius is unfortunately unknown in nature to the
writer, otherwise additional characters would be given to enable
recognition of this very poorly characterized genus, the diagnosis
of which fails to mention the chief characters usually employed
for this purpose, such as the structure of palpi, and gives only a
very ambiguous and contradictory account of the eyes and an
inefficient description of the legs and the tarsi.

Genus Cephaloclerus Kuwert

C ephaloclerus Kuwert, Ann. Soc. Ent. Belg., XXXVII, 1893,
p. 486. — Kraatz, Deutsch. Ent. Zeit., 1899, p. 105. — Schenkling,

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133

Bull. Mus. d’Hist. Paris, VIII, 1902, p. 330 (in species). —
Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 90. — Hintz,
Deutsche. Ent. Zeitschr., 1905, p. 312. — Schenkling, Deutseh. Ent.
Zeit., 1906, heft 2, p. 300. — Gahan, Ann. Mag. Nat. Hist., Lond.,
(8) V, 1910, p. 65. — Schenkling, Col. Cat., (Junk) Cleridae,
1910, p. 111.

Hydnocera Boheman, Ins. Caffr., I, fasc. 2, 1851, p. 511.

Diagnosis of Cephaloclerus, revised

Head large, rather strongly rounded, with the eyes much
broader than the thorax, clypeus strongly emarginate, labrum
emarginate; terminal segment of maxillary palpi short, slender,
cylindrical, acute at apex ; that of labial palpi larger, bell-shaped.
Antennae very short, scarcely longer than the thorax, eleven-seg-
mented, slender, segments four to ten very short, serrate, clava
elongate, but little distinct, three-segmented, terminal segment
rounded, its apex obtuse, within subacuminate, eyes large, ovate,
entire, moderately convex, prominent. Thorax nearly one-half
longer than broad, angles strongly rounded, margins acute, pos-
teriorly bisinuate, apex truncate, narrower than the head, sides
before the middle nearly straight then behind the middle ab-
ruptly narrowed, posteriorly narrower, basal margin truncate,
thinly reflexed, near apex and near base a transverse impressed
line, surface slightly convex, sides at apex and base narrowly
constricted. Elytra oblong-quadrate, narrower than the head,
nearly three times as long as the thorax and twice as broad as
the thorax at base, base slightly broadened, obliquely truncate,
humeri strongly rounded, not elevated, sides thinly margined,
straight (sinuate in C. basalis), apex conjointly rounded, coarsely
and deeply punctate. Legs slender, posterior tarsi viewed from
above evidently four-segmented, segments one, two and three
dilated, penultimate segments lamellate, ungues simple.

Genotype. — Cephaloclerus corynetoides Kuwert. Monobasic.

Geographical range, 23 species, all of Africa.

Genus Eurycranium Blanchard

Eurycranium Blanchard, Hist. Ins., II, 1845, p. 87. — Chenu,
Encycl. d’Hist. Nat. Col., II, 1860, p. 245.

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Eurycranus Blanchard, Hist. Ins., II, 1845, p. 84. — Gemm. et
Harold, Cat. Col., VI, 1869, p. 1745 (pars). — Gahan, Ann. Mag.
Nat. Hist., (8) V, 1910, p. 64 and 65.

Eurymetopum Blanchard, Voy. d’Orb., 1844, p. 92.1 — White,
Nomenc. Col. Ins. Brit. Mus. Cler., IV, 1849, p. 42. — Chevrolat,
Cat. des Cler., Rev. Mag. Zool., 1874, p. 275.

Epiclines Spinola, Gay Hist. Chile, Zool., IV, 1849, p. 386
(pars). — Lacordaire, Gen. Col., IV, 1857, p. 463 (pars). — Gem-
minger et Harold, Cat. Col., VI, 1869, p. 1745 (pars). — Gorham,
Cist. Ent., II, 1876, p. 91 (“nil ad rem”). — Lohde, Ent. Zeit.
Stett., IX, 1900, p. 71. — Schenkling, Gen. Ins., (Wytsman) Cler.,
1903, p. 71 (pars). — Schenkling, Col. Cat., (Junk) Cler., 1910,
p. 99 (pars).

Clerus Spinola, Mon. Cler., II, 1844, p. 158 (in species).

Thanasimus Spinola, Gay Hist. Chile, Zool., IV, 1849, p. 392
(pars). — Fairmaire et Germain, Col. Chilensia, Paris, II, 1861,
p. 3. — Philippi, Stett. Ent. Zeit., XXV, 1864, p. 266 (pars). —
Philippi, Ann. Univ. Chile, Santiago, XXVI, 1865, p. 656 (in
species).

Dereutes Chevrolat, Mem. des Clerites, 1876, p. 29 (pars) (a
group name only!).

Thereutes Harold, Col. Hefte, XVI, 1879, p. 248 (pars)
(inutilis emendat.) .

Diagnosis of Eurycranium, revised

Head short, very wide across the eyes, which are very promi-
nent and nearly entire. The maxillary palpi are short, the ter-
minal segment elongate, conico-cylindrical, that of the labial

i Eurymetopum Blanchard is preoccupied by Eurymetopon Eschscholtz,
Atlas, Zoologique du voyage du capitaine Kotzebue, fascicule 2, p. 8, pi. 18,
figs. 1 and 2, to designate a genus of the family Tenebrionidae. Later
Schonherr (Genera et Species Curculionidum, Paris, VI, 1, 1840, p. 112)
employed the name Eurymetopus to designate a genus of Tetramerous coleop-
tera, of the Brachypterous Curculionides. This author has described but one
species ( E . fallax ) of the great number now known from South America.
Chevrolat says (Mem. Cler., p. 30), “The three names represent the same
root. That [of Dereutes ] which I employ indicates the mode of life of the
greater part of the true derides. Not only do they live on insects in the
larval state, but arriving at the perfect state they run on the bark of trees,
pouncing upon those that they meet devouring them with avidity.”

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palpi securiform, very large, the mandibles are rather small with
a feeble tooth on internal side. Antennae slender, long, eleven-
segmented, nearly as long as head and thorax united, the funicle
very long with the three last segments of antennae forming a
small ovate, serrate clava. Prothorax long, cylindrical and con-
stituting nearly a third of the total length of the insect, sides of
thorax more or less broadly dilated at middle, apical and basal
constrictions and transverse impressions usually quite feeble,
basal margin thinly reflexed. Elytra rather long, entirely cover-
ing the abdomen, flexible, suture closed, apex conjointly rounded,
surface very uneven, puncturing imperceptible. Legs moder-
ately long, slender, tarsi five-segmented, first segment very short
and small, but still visible from above, the second longer than
the first but still less stout, third slightly shorter than the second,
fourth dilated.

Genotype. — E. maculatum Blanchard, by present designation.

Geographical range, South and Central America.

Genus Paupris Sharp

Paupris Sharp, Ent. Monthly Mag., XIII, 1877, p. 271. — Gor-
ham, Trans. Ent. Soc. Lond., 1877, p. 263. — Schenkling, Gen. Ins.,
(Wytsman) Cler., 1903, p. 95. — Broun, Man. New Zealand Col.,
I, 1880, p. 331. — Schenkling, Gen. Ins., (Wytsman) Cler., 1903,
p. 95. — Gahan, Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. — Schenk-
ling, Col. Cat., (Junk) Cler., 1910, p. 112.

Diagnosis of Paupris, revised

Elongate, apterous. Head (the eyes included) rather broader
than the thorax at its broadest part. Eyes rather small, moder-
ately prominent, coarsely granulated, with a very small tri-
angular excision. Maxillary palpi with terminal segment small
and cylindrical, not at all dilated, while that of the labial palpi
is very large, transversely broad, and dilated. Antennse short and
stout, inserted well before the eyes, eleven-segmented, segment
two shorter than the third, segments of the funicle three to eight
all subequal in length, segments nine to eleven each broader than
long, about twice as broad as the preceding segments, the terminal
segment as broad as long, its apex subacuminate, these segments

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forming a rather lax clava. Prothorax elongate and narrow,
longer than broad, sides at middle somewhat dilated, but scarcely
tuberculate. Elytra small and narrow, much abbreviated,
usually nearly three dorsal segments shorter than the abdomen,
base narrow, humeri absent, suture closed, elongate-obovate, very
sparsely covered with short erect coarse hairs, apex conjointly
rounded. Legs long and rather stout, tarsi moderately broad,
five-segmented, first segment very small and short, nearly covered
by the elongate second segment, segments three and four each
shorter than the second, and strongly lamellate, ungues small,
simple.

Genotype. — Paupris aptera Sharp. Monobasic.

Geographical range, one species of New Zealand.

Genus Isolemidia Gorham

Isolemidia Gorham, Trans. Ent. Soc. Lond., 1877, pp. 249,
257. — Gorham, Biol. Centr.-Amer., Ill, pt. 2, 1883, p. 177. —
Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 92. — Schenk-
ling, Col. Cat., (Junk) Cler., 1910, p. 107. — Gahan, Ann. Mag.
Nat. Hist., (8) V, 1910, p. 65. — Wolcott, Publ. Field Mus. Nat.
Hist., Zool., VII, 1910, p. 376 (in species). — Chapin, Proc. Ent.
Soc. Wash., XXII, 1920, pp. 51 and 52 (in species).

Diagnosis of Isolemidia , revised

Head with the eyes broader than the thorax, eyes very large,
globular, entire or at most minutely emarginate, maxillary palpi
subfiliform, terminal segment truncate at apex ; that of the labial
palpi securiform. Thorax subcylindrical, but often short and
sometimes quadrate or broader than long, the sides broadly
rounded or tuberculate at middle, strongly constricted near apex
and base, the basal and apical transverse impressed lines very
distinct. Antennae very short, usually little longer than the
head, as in Lemidia, eleven-segmented, those segments succeed-
ing the basal segment a little longer than wide, the three last
transverse, forming a small, short, rather connate, oblong, clava.
Elytra elongate, usually much broader than the thorax, and
broadest behind the middle, the apex conjointly rounded, usually
covering the abdomen, roundly truncate ( pulchella Gorh.), the

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humeri distinct, sometimes with a very small callus (/. cariniceps
Wole., the elytra are much abbreviated, the suture widely dehis-
cent, the apices separately obtusely rounded with a few well
developed teeth). Legs long and slender, posterior femora mod-
erately long, tarsi five-segmented, the basal segment as in Lemidia,
atrophied, second segment covering the first, ungues simple.

Genotype. — I. pulchella Gorham, designated by Gorham.

Geographical range, thirteen species of South and Central
America.

Genus Emmepus Motschoulsky

Emmepus Motschoulsky, Bull. Moscou., XVIII, 1845, p. 41 (sub
Staphylin.). — Lacordaire, Gen. Col., IV, 1857, p. 494. — Chenu,
Encycl. d’Hist. Nat. Col., II, 1860, pp. 82 and 266 (sub Staphy-
lin.).— Gorham, Trans. Ent. Soc. Lond., 1877, p. 263. — Reitter,
Verh. Nat. Ver. Brunn., XXXII, 1893 (1894), p. 39. — Reitter,
Best. Tab., 1894, p. 6.— Reitter, Wien. Ent. Zeit., XV, 1896, p.
283. — Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 96. —
Gahan, Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. — Schenkling,
Col. Cat., (Junk) Cler., 1910, p. 112.

Brachyclerus Fairmaire, Compt. rend. Soc. Ent. Belg., XXVII,
1883, p. 157. — Bedel, Ann. Soc. Ent. Fr., (6) VII, 1887, p. 197.

Diagnosis of Emmepus, revised

Body slender, small. Head transversely quadrate, front broad,
mandibles internally with a distinct tooth near apex. Maxillary
palpi with the terminal segment small, nearly cylindrical, at apex
much smaller ; that of labial palpi large, securiform. Eyes large,
prominent, finely granulated, ? entire. Antennae very short,
eleven-segmented, the two first segments large and thick, the
succeeding segments much shorter and more slender, the two
terminal segments forming a ball-like clava, first segment of
clava (the tenth of the antennae) much swollen, partly covering
the very small apical segment. Prothorax as long as broad, the
sides at middle broadly subangulately dilated. Elytra twice as
long as broad, much larger than the thorax, much shorter than
the abdomen, suture strongly dehiscent, scarcely covering the
abdomen behind the middle. The wings long, extending well
beyond the apex of the abdomen which it covers. Legs long and

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slender, tarsi rather long and slender, distinctly five-segmented,
first segment longer than the second, segments three and four
each successively shorter than the preceding segments, segments
one to four with lamellae beneath, segment five elongate-clavate,
much longer than the two preceding segments united, ungues
simple.

Genotype. — Emmepus arundinis Motsch. Monobasic.

Geographical range, three species known from Caucasus, 'Cas-
pian Sea, South Algeria, Turkestan and Ceylon.

Genus Theano Castelnau

Theano Castelnau, Silberm. Rev., IV, 1836, p. 51. — Lacordaire,
Gen. Col, IV, 1857, p. 493.— Chenu, Encycl. d’Hist. Nat. Col, II,
1860, p. 268. — Gorham, Trans. Ent. Soc. Lond, 1877, p. 363. —
Schenkling, Gen. Ins, (Wytsman) Cler, 1903, p. 97. — Gahan,
Ann. Mag. Nat. Hist, (8) V, 1910, p. 65. — Schenkling, Col. Cat,
(Junk) Cler, 1910, p. 113.

Diagnosis of Theano , revised

Body small, rather long. Head with the eyes broader than the
thorax. Labrum entire, its front arcuate, mandibles with an
internal tooth near apex. Terminal segment of labial palpi large,
securiform, acuminate at apex. Eyes moderately large, promi-
nent, entire. Antennae short, slender, eleven-segmented, the first
two segments equal, rather large, the third segment slender, the
succeeding five segments rather slender, trianguler, the last three
segments forming a swollen, ovate or somewhat globular clava.
Thorax much narrower than the head, rounded at the sides.
Elytra somewhat elongate, surface very coarsely punctate. Legs
long and slender, posterior femora extending slightly beyond the
apex of the abdomen, tarsi with the basal segment conical, the
succeeding three segments short, dilated and bifid, provided
with prolonged lamellae beneath.

Genotype. — Theano pusilla Cast. Monobasic.

Geographical range, one species from South America (Colom-
bia).

Subgenus Callimerus Gorham

Callimerus Gorham, Cist. Ent, II, 1876 (1875-1882), p. 65. —
Schenkling, Gen. Ins, (Wytsman) Cler, fasc. 13, 1903, p. 24. —

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139

Gahan, Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. — Schenkling,
Col. Cat., (Jnnk) Cler., 1910, p. 26. — Chapin, Philipp. Jonr. Sci.,
XXV, No. 2, 1924, pp. 180 and 191. — Pic, Exot.-Ent., fasc. 54,
1929, p. 17 (pars).

Xylobius White, Nomenc. Col. Brit. Mus., Cler., IV, 1849, p. 50
(in species) .—Westwood, Proc. Zool. Soc. Lond., XX, 1852, p. 40
(in species).

Lemidia Thomson, Mus. Scient., II, 1860, p. 61 (in species). —
Pascoe, Jour. Ent., I, 1860, p. 48. — Chevrolat, Rev. Mag. Zool.,
1874, p. 321. — Kuwert, Ann. Soc. Ent. Belg., XXXVII, 1893, p.
484.

Caloclerus Kuwert, Ann. Soc. Ent. Belg., XXXVII, 1893, p.
480. — Schenkling, Ann. Soc. Ent, Belg., VL, 1901, p. 105.

Diagnosis of Callimerus, revised

Form elongate, or oblong, head with eyes slightly broader than
apex of thorax, eyes prominent, finely granulate, very slightly
emarginate at base of antennae. Labrum entire ; mandibles stout,
falcate, with internal tooth near apex ; terminal segment of maxil-
lary palpi twice as long as preceding segment, subulate ; that of
the labial palpi elongate-triangular. Antennae rather short,
slender, distinctly eleven-segmented, first segment stout, bent, at
least twice the length of the second, which is from nearly as long
as broad to equilateral and globular, segments three to eight much
longer than broad, each segment shorter than the preceding, ninth
to eleventh forming a lax, oval clava, the eleventh oval at apex.
Thorax longer than broad, somewhat constricted before and be-
hind, moderately to strongly dilated at or before the middle,
anterior coxal cavities very widely open behind. Elytra entirely
covering the abdomen, long, narrow, sides parallel or gradually
becoming narrower posteriorly, adorned with white or yellow
scales, these scales often forming a, definite pattern, apex some-
times abbreviated and truncate or rarely each mucronate, surface
variously punctate. Abdomen with six visible ventral segments ;
in the male the terminal segments are often profoundly modified ;
in the female the modification, if any, occurs on the penultimate
segment. Legs rather long, posterior tibiae with or without a
notch with comb of hairs near its apex; tarsi moderately long,

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five-segmented, the basal segment the longest. Ungues nearly
simple or with a broad plate-like tooth.

Subgenotype. — Clerus ( Xylobius ) dulcis Westw., designated
by Gorham.

Geographical range, 102 species known from Indo-Malaysia.

Subgenus Cucujocallimerus Pic

Cucujocallimerus s. g. Pic, Mel. Exot.-ent., fasc. LIV, 1929,
pp. 17-18.

Callimerus auct. (pars).

Diagnosis of Cucujocallimerus, slightly revised

Prothorax less than, or little longer than broad, or sometimes
very elongate, posterior margin less narrow, often rather broad,
or sometimes indistinct, femora more or less slender or a little
stouter, sometimes the posterior stouter than the others; the
tarsi the most often slender and long. Form of the body oblongo-
elongate or narrow. Antennas less short. Head with the eyes
much broader than the front part of the prothorax, this very im-
pressive. Body wholly, as a matter of fact, depressed above.

“The sub-genus Cucujocallimerus mihi has the legs principally
the posterior femora much longer than the others, whereas the
greater portion of the species of the s. genus Callimerus has them
little or moderately longer than the others” (Pic).

Subgenotype. — Cucujocallimerus ( Callimerus ) coomani Pic.,
designated by Pic.

Geographical range, 1 species of Tonkin.

Subgenus Brachycallimerus Chapin

Br achy callimerus Chapin, Philipp. Jour. Sci., XXV, No. 2,
1924, pp. 180 et 190. — Corporaal, Eev. Fran^aise d’Ent., IV,
1937, p. 60.

Callimerus auct. (pars), loc. cit., p. 190.

Crassocallimerus Pic, Mel. Exot.-ent., LIV, 1929, pp. 17 et 18.

Diagnosis of Brachycallimerus, only slightly revised

Head broad, eyes very prominent, very slightly emarginate
near antennal socket; labrum entire, mandibles broad and flat,

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141

falcate, with a sharp tooth internally near apex; antennae short
and compact, eleven-segmented, first segment short and stont,
second nearly spherical, third to sixth longer than broad, seventh
to eleventh broader than long, each broader than the preceding,
together forming a compact obtriangnlar five-segmented clava.
Thorax broader than long, polished, with a few distinct punc-
tures. Elytra broader at base than the thorax, punctures numer-
ous, not seriate. Entire dorsal surface of insect devoid of scales.
Abdomen with six visible ventral segments, secondary sexual
modifications conspicuous. Legs moderate in length, posterior
tibiae with subapical notch, tarsi rather long and slender, five-
segmented, first segment longer than the second, distinct and not
covered by the second, ungues broadly toothed at base.

Subgenotype. — Callimerus latifrons Gorh., designated by
Chapin.

Geographical range, Indo-Malaysia.

Chapin states that “The insects which I include in this new
genus have heretofore been classified with the species of Calli-
merus Gorh. They differ from those species in their broad and
compact form, short and compact antennae, and the total absence
of scales from the upper surface. In addition to the genotype,
I would include the following species : latesignatus Gorh., rus-
ticus Gorh., pectoralis Schklg., and probably trifasciatus Schklg.”
Corporaal (1937 : 60) includes the last named species without
question and adds pallidus Gorh., and doesburgi Corp., n. sp., as
members of this genus.

Genus Evenus Castelnau

Evenus Castelnau, Silb. Rev., IV, 1836, p. 41. — Spinola, Rev.
ZooL, 1841, p. 75. — Klug, Abhandl. Berl. Akad. 1842, p. 315. —
Spinola, Mon. Clerites, II, 1844, p. 28. — Lacordaire, Gen. Col.,
1857, p. 469. — Chenu, Encycl. d’Hist. Nat. Col., II, 1860, p. 266.
— Gorham, Trans. Ent. Soc. Lond., 1877, p. 249. — Schenkling,
Gen. Ins., (Wytsman) Cler., 1903, p. 89. — Gahan, Ann. Mag. Nat.
Hist., (8) V, 1910, p. 65. — Schenkling, Deutsche Ent. Zeitschr.,
1906, p. 300. — Schenkling, Col. Cat., (Junk) Cler., 1910, p. 111.

Diagnosis of Evenus, revised

Body very narrow, filiform. Head large, constricted behind,
front concave. Maxillary palpi small, terminal segment cylin-

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drical, rounded at apex ; labial palpi long, three times as long as
the maxillary palpi, terminal segment elongate, securiform,
slender at base. Eyes large, very prominent transversely oval,
not visibly emarginate. Antennae very short, inserted between
the eyes and very near their extreme anterior margin, eleven-
segmented, first segment rather stout, long, cylindrical, segments
three to eight obconical, small but distinct, each progressively
broader but without becoming longer, the ninth as broad as the
eighth and forming with the two last segments a narrow, elongate
clava, terminating in an acute point, a little flattened and at their
articulations rather serrate rendering at least very doubtful
the independent mobility of the tenth and eleventh segments.
Thorax narrow and elongate, disk depressed and narrowed in
front, sides feebly dilated, front and hind margins equal in width,
anterior coxal cavities open behind. Elytra depressed, much
broader than base of thorax, very elongate parallel, but not reach-
ing to apex of the abdomen, suture closed, irregularly rounded
at apex, surface rather smooth (in the typical species sometimes
seriately punctate). Legs elongate, thin, the posterior pair
twice as long as the others, their femora reaching beyond the apex
of the elytra, tibiae rather longer than the femora, straight and
cylindrical, . tarsi very long, five-segmented, equally visible, long
and narrow, the posterior having the two first elongate conical,
the first shorter than the second by about one-third, third and
fourth short, bifid, lamellate beneath, the second longer than any
of the others, subequal to segments three and four united, and
somewhat similarly depressed, a little dilated and feebly emargi-
nate at apex, the fifth segment shorter than either of the preced-
ing two, terminated by two large and short simple ungues, all
tarsi pubescent beneath.

Genotype. — Evenus filiformis Cast. Monobasic.

Geographical range, thirteen species all of Madagascar.

Genus Lasiocallimerus Corporaal
Lasiocallimerus Corporaal, Tijds. v. Ent., 82, 1939, p. 194.

Diagnosis of Lasiocallimerus Corporaal
il Tarsorum articulo primo longiore quam secundo. Statura
generi Brachycallimero similis , sed antennce 10-articulatce, arti-

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143

culo robusto, articulo secundo ceque crasso, paulo breviore,
articulo tertio subelongato, ariiculis 4-9 transversis, articido
ultimo subelongato, ovato, ad apicem subacuminato. Corpus
longe pilosum et, in thorace, dense squamuloso-villosum. Pedes
quotes in genere Brachycallimero, tibiis sine tuber culo, tar sis
latis, unguiculis latis, ad apicem emarginatis.

“With the first tarsal segment longer than the second. Head
broad; eyes [rather large] very prominent, [finely facetted, with
long, erect, pale hairs] very slightly emarginate near antennal
socket, labrum entire, mandibles broad and flat, falcate. An-
tennae short and compact, ten-segmented ; first segment short and
stout, second of equal thickness, but shorter, third a little longer
than broad, fourth to ninth broader than long, tenth longer than
broad, ovate, a little acuminate. Prothorax of the same form as
in Brachycallimerus, heavily punctate, under the longer pilosity
closely covered with crisp, scale-like hairs. Elytra broader than
prothorax, evenly punctured, but not in rows. Abdomen with
six visible ventral segments. Legs of moderate length, tibiae
without notch, tarsi short and broad, claws broad, emarginate at
tip” (Corporaal).

Genotype. — Lasiocallimerus vestitus Corp., designated by Cor-
poraal.

Geographical range, one species of Java.

Genus Allelidea Waterhouse

Allelidea Waterhouse, Trans. Ent. Soc. Lond., II, 1839, p. 193.
— Lacordaire, Gen. Col., IV, 1857, p. 473. — Chenu, Encycl.
d’Hist. Nat., Col., II, 1860, p. 266. — Gorham, Trans. Ent. Soc.
Lond., 1877, p. 263. — Blackburn, Trans. Roy. Soc. South Austral.,
XIV, 1891, p. 302. — Schenkling, Gen. Ins., (Wytsman) Cler.,
1903, p. 97. — Gahan, Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. —
Schenkling, Col. Cat., (Junk) Cler., 1910, p. 112.

Diagnosis of Allelidea, revised

Body elongate, cylindrical. Head with the eyes as broad or
broader than the thorax, constricted posteriorly, front verticle,
labrum transverse, emarginate. Terminal segment of maxillary
palpi ?, that of labial palpi oblique, securiform. Mandibles in-

144

Journal New York Entomological Society

[Vol. LII

ternally with a small tooth before the apex. Eyes moderately
prominent, entire. Antennae very short, eleven-segmented, ar-
ticulations very distinct, segments very small, gradually growing
larger, the three last segments slightly larger than the others
forming a very lax elongate-oblong clava. Thorax longer than
broad, narrow and elongate, subcylindrical, sides a little before
the middle roundly dilated, at bast strongly narrowed. Elytra
not much broader than thorax at base, seriately punctured, elon-
gate, gradually and very feebly broader posteriorly, apex serrate,
as long as the abdomen (J' ?), or much shorter (5 ?). The three
pairs of legs equal in length, long and slender, posterior femora
slightly shorter than the abdomen, tarsi five-segmented, first seg-
ment not much longer than the succeeding segments together,
segments two to four with bilobed membraneous appendices, fifth
segment moderate. Ungues with membraneous appendices.

This is a small insect with the facies of Phyllohoenus.

Genotype. — Allelidea ctenostomoides Waterh., designated by
Gorham.

Geographical range, seven species of Australia.

Remarks on genera more or less allied to the genera of the
subfamily Phyllobseninae, or transferable to that subfamily.

Genus Brachyptevenus Pic

Brachyptevenus Pic, Mel. Exot.-ent., LXXI, 1939, pp. 24-25.

Diagnosis of Brachyptevenus , revised

Head broader than the thorax, posteriorly long attenuate, not
strangulate. Thorax elongate, sides sinuate, middle impressed
posteriorly. Elytra but little reduced, anteriorly subattenuate,
narrow and elongate, apex slightly broader. Legs slender, ab-
domen distinctly exceeding the elytral apex. This new genus is
near E venus Cast. It is clearly characterized by the form of the
head and the elytra. It is established for the following new
species B. niger Pic.

Genotype. — Brachyptevenus niger Pic. Monobasic.

Geographical range, 1 species of Madagascar.

The writer has been compelled by reason of the brief and un-
satisfactory diagnosis of this genus to omit it from consideration
in the body of this account.

June, 1944]

Wolcott: Phyllob^enin^e

145

Genus Achlamys Waterhouse

Achlamys Waterhouse, Cist. Ent., II, 1875-1882 (1879), p.
530. — -Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 16. —
Schenkling, Col. Cat., (Junk) Cler., 1910, p. 17.

Diagnosis of Achlamys , revised

Form cylindrical. Head (with the eyes) broader than the
thorax, the eyes prominent, coarsely granulate, scarcely emargi-
nate in front. Antennae as long as the head and thorax taken
together, the first segment thick, ovate, the second segment small,
the third elongate, subcylindrical, the fourth, fifth and sixth be-
coming gradually stouter and shorter, the seventh as broad as
long, the eighth, ninth, tenth and eleventh segments forming a
distinct four-segmented clava. Apical segment of the maxillary
palpi elongate, a little narrowed before the apex ; apical segment
of the labial palpi securiform. Prothorax very convex, one-third
longer than broad, very slightly constricted before the front
margin, strongly constricted before the base. Elytra parallel,
a little broader than the thorax, each with seven lines of deep
oblong punctures which do not extend quite to the apex, which
is smooth and obtuse. Tarsi with five distinct segments, the
ungues with a single tooth at the base.

“Closely allied to Pallensis, but with prominent eyes and dis-
tinct club to the antennas” (Waterhouse).

The writer believes Achlamys should be removed from Tillinae
to the Phyllobaeninae and there with Paupris Sharp, both genera
having coarsely facetted eyes, a character exceptional in this sub-
family, these two genera may be placed as a distinct group or
tribe designated by the term Pauprini.

Gahan (1910: 65) has transferred the genus Ellipotema
Spinola to the subfamily Corynetinae, and the genera Cleropiestus,
Calendyma Lac., and Epiclines Chevr., to the subfamily Clerinas,
he also briefly differentiates ( loc . cit p. 64) the genus Epiclines
and Eurycranus (= E-ury cranium Blanch.) and calls attention to
the long-standing and absolutely absurd erroneous synonymy
which has been perpetuated ever since the issuance of Lacor-
daire ’s Genera des Coleopteres. Of the species in our Catalogues
under Epiclines only a few, as far as can now be determined,

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Journal New York Entomological Society

[Vol. LII

really belong to that genus, those that should be so placed are
as follows: gayi Chevr., (nec Spin.), cenea Phil., basalis Blanch.,
puncticollis Spin., ruficollis Phil., similis Schklg., advena Chevr.,
viridis Phil., and probably tristis Spinola. The rest of the listed
species with scarcely an exception should go to the genus Eury-
cranium Blanchard.

Genotype of Epiclines, E. gayi, Chevrolat. Monobasic.

Geographical range, Chile.

The genera of this subfamily may be grouped according to
natural characters as follows:

Group 1

Clava two-segmented; basal segment of hind tarsi short, cov-
ered above by the second segment, prothorax variously propor-
tioned.

Pliyllobcenus Dejean, Lemidia Spinola, Isohydnocera Chapin.

Group 2

Clava indefinitely two- or three-segmented; basal segment of
hind tarsi short, covered above by the second segment, prothorax
with length and breadth subequal.

Wolcottia Chapin, Parmius Sharp, Neohydnus Gorham,
Metaxina Broun.

Group 3

Clava three-segmented ; basal segment of hind tarsi short, cov-
ered above by the second segment, prothorax with length usually
greater than breadth.

Abrosius Fairmaire, C ephaloclerus Kuwert, Eury cranium
Blanchard, Paupris Sharp, Isolemidia Gorham.

Group 4

Clava two-segmented • basal segment of hind tarsi long, not cov-
ered by the second segment, prothorax as long as broad.

Emmepus Motchoulsky.

Group 5

Clava three-segmented; basal segment of hind tarsi long, not
covered by the second segment, prothorax at least as long as
broad.

June, 1944]

Wolcott: Phyllob^nin^

147

Theano Castelnau, sub. gen. Callimerus Gorham, sub. gen.
Cucu jo callimerus Pic.

Group 6

Clava five-segmented ; basal segment of hind tarsi long, not cov-
ered by the second segment, prothorax broader than long.

Sub. gen. Br achy callimerus Chapin.

Group 7

Clava indistinctly four-segmented ; basal segment of hind tarsi
long, not covered by the second segment, prothorax longer than
broad.

Evenus Castelnau.

Group 8

Clava one-segmented ; basal segment of hind tarsi long, not cov-
ered by the second segment, prothorax broader than long.

Lasio callimerus Corporaal.

Group 9

Clava indistinctly three-segmented ; basal segment of hind tarsi
long, prothorax longer than broad.

Allelidea Waterhouse.

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Schneirla: Insect Behavior

153

THE REPRODUCTIVE FUNCTIONS OF THE ARMY-
ANT QUEEN AS PACE-MAKERS OF THE
GROUP BEHAVIOR PATTERN

T. C. Schneirla1

Although they are among the commonest ants encountered in
the tropical forests of the Old and New Worlds, the dorylines have
long remained among the least known. In Africa and Asia the
“driver ants’ ’ and in tropical America the “legionary” or
“army” ants of this subfamily present many intriguing problems,
especially in the predatory raids and nomadic life of their colonies
and the biological status of their huge wingless fertile females, the
dichthadiigynes. These last individuals are not very well known
to science, since up to a relatively short time ago virtually no pre-
cise information was available concerning their relations to the
colony. Actually we find, beyond propagative functions which
differ rather strikingly in some respects from those of fertile
females in other insects, the properties of the doryline queen make
her a factor of critical importance in the general process of colony
behavior.

This paper is directed toward accounting for the essential onto-
genetic basis of the behavior pattern characteristic in the subgenus
Eciton s. str., an American branch of the subfamily Dorylinae.
Our study specifically concerns these terrestrial army ants, espe-
cially the type species of the subgenus, E. hamatum. The results
may prove eventually to have relevance among the dorylines far
beyond the conditions of the species on which this report is based.2

1 Department of Animal Behavior, American Museum of Natural History,
New York, and Department of Psychology (Washington Square College),
New York University.

2 Acknowledgments : The field work for this study was subsidized by a
grant from the Bache Fund of the National Academy of Sciences. Prepara-
tion of the manuscript was made possible by a grant from the Committee for
Research in Problems of Sex (National Research Council), administered by
Dr. Frank A. Beach. The writer also wishes to express his thanks to Mr.
James Zetek, Custodian of the Barro Colorado Island Biological Reservation,
for his hospitality and his unstinting help with arrangements for work at the
station.

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Journal New York Entomological Society

[Vol. LII

The chief characteristics of the behavior pattern of E. hamatum
involve the formation of bivouacs, raiding, and bivouac-change
movements which in series may be termed migrations. In this
species the temporary nest or bivouac of a given colony is a more or
less cylindrical mass formed by the clustered bodies of ants alone,
without foreign materials, usually depending from a log or vines,
or hanging against a tree. From his studies in the rainy season
the writer (1933; 1938) has described two distinctive conditions
through which colonies of E. hamatum pass successively in alter-
nation at such times : the nomadic and the statary activity phases.
Roughly, each phase lasts close to 20 days before it is succeeded
by the other. The principal characteristics of the nomadic phase
are 1) highly developed daily raids and 2) a regular change of the
bivouac site at the termination of each day. In rather sharp con-
trast, the statary phase is marked by 1) less developed raids and
2) occupancy of the same bivouac site throughout the interval.
In addition there are other features which characterize each of
these behavior phases. These are essentially intrinsic to the
colony, yet as we shall find they are intimately involved in the
entire system of events in Eciton behavior.

Our essential hypothesis is that a close relationship exists be-
tween behavior outside the bivouac (i.e., raiding and bivouac-
change processes) and conditions within the bivouac, and that this
pattern of events basically depends upon the functions of the
reproductive agent of the colony. Accordingly in this paper at-
tention focusses upon the role of the mother queen, whose proper-
ties apparently are of key importance in the entire mechanism
of the Eciton behavior pattern.

Material for this study was gathered and most of the field ob-
servations were carried out on Barro Colorado Island in the
Panama Canal Zone, in the rainy season months of 1936 and 1938
(May to September). In that locality the rainy period usually
begins in April and ends in early December. Investigations of
the Eciton problem have yet to be conducted in the dry season.
This limitation is important, since it is very probable that rainy-
season conditions are optimal for the appearance of relationships
such as we shall describe. In an indirect manner the results of
this study suggest that with extreme changes in prevalent atmos-

June, 1944]

Schneirla : ' Insect Behavior

155

pheric conditions important variations may occur in the described
behavior pattern. In view of this, it is a fact of some importance
that onr evidence was gathered in a portion of the Caribbean
lower rain-forest zone in which the rainy and dry seasons are dif-
ferentiated rather sharply from year to year. What differences
will be found under more variable meteorological conditions rep-
resent an interesting problem.

PROPERTIES OF THE ECITON QUEEN

From the time Andre (1885) captured the first dichthadiigyne
from the subterranean bivouac of an E. ( Labidus ) coecum colony
the list of captures has grown very slowly, and although fertile
females are at present known from nearly 25 of the more than 100
recorded species of Eciton and all of the eight recorded species of
Eciton s. str. (Bruch, 1934), they are still among the most highly
prized collector ’s items. In view of the great difficulty of captur-
ing them, it is scarcely surprising that these rare insects have been
almost invariably clapped into alcohol when taken, under the
influence of what Creighton has appropriately termed ‘ ‘ collector ’s
itch. ’ ’ This practice of course has not precluded the accumulation
of much valuable information concerning the external morphol-
ogy and the taxonomic affinities of the specimens; yet it has
unfortunately kept in the remote background several important
questions which can find their answers only in study of the living
queen.

Thus it is unfortunate that from the records of more than a
score of collections very little can be learned about the behavior
and biological properties of the living individual. With the ex-
ception of general observations carried out by Wheeler (1900)
with captive colonies of E. schmitti in Texas, no special investi-
gations have been attempted. The present study stems from a
general investigation of army-ant behavior which has revealed the
probability that the dichthadiigyne plays a crucial though in-
direct role in the phenomena of Eciton behavior (Schneirla,
1934; 1938).

Eciton queens may be found in either of two very different con-
ditions, the “normal” or contracted condition and the physo-
gastric or egg-producing condition (see Fig. 1). For the present
study, the dichthadiigyne material comprised 13 queens of E .

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hamatum and one of E. burchelli in the contracted condition, and
one queen of E. hamatum in the physogastric condition. In each
case, field notes on the colony from which a given queen was
taken covered the general situation of the colony and its behavior
(especially its raids and bivouac-change movements) over a period
of days, the status of the brood or broods and the condition of the
queen at the time of capture. Each queen was kept alive in the
laboratory as long as possible after capture, for observation and
test both alone and in relation to the workers of her colony.

There is a notable disparity in our material as to condition of
the queen when captured. Why so many of the cases were in the
contracted condition and only one case physogastric when taken
will become clear as the general circumstances of Eciton life are
disclosed.3 For a number of reasons, physogastric army-ant
queens are exceedingly difficult to capture. There is only one
other case on record in which such an individual was observed
during her short span of life in captivity, that of a physogastric
queen of E. ( Labidus ) coecum taken by Weber (1941) in Trini-
dad. Fortunately we are not forced to depend upon direct evi-
dence concerning the queen at all stages, since an abundance of
other evidence coordinates nicely with results derived from work
on the queen herself.4

A descriptive contrast of queens and workers. — The Eciton
queen (Fig. 1) stands out as strikingly unique among fertile in-
sect individuals and as a highly distinctive member of her colony.
We may use the queen of E. hamatum as example.5 Unlike the
queens of virtually all other ants, she is wingless throughout life.
In color she is almost uniformly ferrugineous mahogany, darker
than the yellowish brown which characterizes the workers, and
lacks the cephalic paleness of major workers. She differs strik-

3 It is a striking fact that among 16 captured queens representing ten
Eciton s. str. species and sub-species, reported by various authors prior to
1942 (see e.g., Wheeler, 1921; Bruch, 1934), all were in the contracted
condition.

4 In the present paper the supporting evidence is reported rather concisely.
Further details and additional evidence will be incorporated in a monograph
on this subject, now in preparation with the collaboration of Dr. Harold R.
Hagan of the Department of Biology, City College of New York.

s The reader is referred to Wheeler Js systematic descriptions of the first
queens of E. hamatum (1925) and E. hurchelli (1921) to be discovered.

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ingly from the workers in gross size and in anatomical detail.
The greater bulk of the queen is suggested by the fact that her
over-all length (contracted) is close to 17.1 mm. (Av. 6 speci-
mens), whereas the range of body length in the workers is 2.5 mm.
(minor) to 9.5 mm. (major). The polymorphic worker types re-
semble the queen only very broadly, for example both queen and
workers possess well-developed and dentate tarsal claws, near the
inferior occipital corners of the queen’s head there are small
obtuse projections homologous to the acute spines of the worker
major, and the queen possesses strikingly prominent pairs of
epinotal and petiolar horns contrasting with two simple aligned
nodes in the worker. Among many impressive morphological
differences are the large lateral pseudo-ocelli (Werringloer, 1932)
of the queen and the relatively tiny ones of workers, the great
bulk of the queen’s thorax and her well-developed legs as con-
trasted with corresponding slenderness in the worker ’s structures,
and in particular the great size of the queen ’s abdomen contrasted
with the much smaller gaster of the worker. Associated with an
enormous expansion of reproductive functions in the queen and
the probable absence of such functions in the worker, the hamatum
queen’s gaster even when contracted measures near 9.1 mm. in
length and 5.5 mm. in its greatest width, whereas the gaster of the
largest major worker measures only 2.6 mm. in length and 1.8 mm.
in width. The queen’s gaster terminates distinctively in a large
triangular shelf, the hypogynium. Another striking character-
istic of the queen is the marked development of her external
respiratory orifices or stigmata, suggesting a high stage of de-
velopment in the respiratory system generally.

This general morphological comparison may suffice to illustrate
the extent to which the Eciton queen diverges from the workers,
paralleling a functional specialization which correspondingly
differs markedly from that of the workers. The nature and
extent of the dichthadiigyne ’s specialization becomes clear when
we consider some typical characteristics of her behavior and her
place in the life economy of the colony.

Behavior and functional 'properties of the “normal” or con-
tracted queen. — In E. hamatum and probably in Eciton s. str.
generally the reproductive capacity of any given colony is cen-

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tered in a single individual, a fecund dichthadiigyne. A selective
mechanism of some kind evidently is involved which excludes ad-
ditional queens, possibly at a time shortly before or after young
queens are fertilized. Circumstances indicate that this one
fecund individual must pass around nine-tenths of her days in
the rainy season in the resting or contracted condition, which as a
consequence may be termed her “normal” state. Ordinarily she

Fig. 1. Queens of E. liamatum in the contracted (upper) and the physo-
gastric or gravid condition. ( Cf . Fig. 2 and Table I.) Total length of the
contracted queen, 17 mm. (Drawing by Miss Janet Roembild, from photo-
graphs by Dr. Virgil Argo, Dept, of Biology, City College of New York.)

does not leave the confines of the bivouac, except when she is
drawn into a bivouac-change movement, an event that scarcely
ever engages her before nightfall.

Circumstances indicate that the sequestration of the queen may
be due largely to the effect which light exerts upon her. In labo-
ratory tests she reacts specifically to directionalized bright light

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by turning away from the source and moving energetically about
until in darkness. Whenever she chances into the light, facing
toward the source, the queen volte faces abruptly and runs off.
(Among numerous observations, this test was repeated in one
instance 12 times in 30 minutes with the same queen, — always
with the described result.) When illumination is not intense
the response is more variable, yet even then as a rule the queen
settles down only when in a fairly dark part of the nest.

In contrast to the queen, the workers seem virtually incapable of
specific orienting responses to light. They are aroused to activity
by light after a period of darkness; thus colonies are regularly
stimulated to begin their daily raiding after dawn (Schneirla,
1938; 1940). This however is a mere photokinetic effect ( i.e
a generalized excitation by light), and they appear incapable
of orienting to directionalized illumination except in a very crude
fashion as an outcome of restless movements over a considerable
time. That the basis of this outstanding difference between queen
and workers rests in different optoneural equipment is suggested
by the fact that in E. hamatum the gross size of the queen ’s lateral
pseudo-ocellus is more than twice that of the major worker’s.
We should expect to find correspondingly accentuated differences
in the internal structure of the visual receptor, and in related
mechanisms, underlying the queen’s pronounced normal reac-
tivity to light.6

This recalls the fact that the fertilized queens of ants in general
are repelled by light. Whether the Bciton queen, like any others,
is positive to light prior to fertilization cannot be said at present ;
however, the sharpness of the functional queen’s avoidance re-
sponse is clear. The basis of a highly adaptive circumstance is
thereby provided — that the queen is cloistered in a well-protected
situation except when the colony shifts its home site. The prev-
alence of this condition is attested by the fact that Eciton queens
are never seen at other times, as during raids — its effectiveness is
emphasized by the difficulties one experiences in capturing these
queens.

6 Although no queen material was available for Werringloer *s (1932) valu-
able histological study of Eciton visual mechanisms, species differences were
found in optic histology which appear to parallel typical behavior differences.

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That the queen keeps to the bivouac in the daytime, during
raids, is not attributable to any inability to follow the colony’s
trails. In the laboratory, captive hamatum queens may run for
hours at a time in a column following a circular path established
by workers or, when one is set down near a vacated trail of her
own colony in the forest, she follows the route successfully once
she hits upon it. Evidently it is her “photophobic” proclivities
that mainly account for the queen’s sequestration in the bivouac.

Although for a long time the idea has held sway that the army-
ant queen is moved passively about by the workers when the colony
shifts its site, in Eciton s. str. the facts are contrary to this sup-
position. In laboratory nests the queen readily makes her way
about despite the usual added burden of several workers hitch-hik-
ing and dragging from her legs or gaster. The physical char-
acteristics of her well-developed legs give every indication of great
strength, as does her robust frame in general — an impression
borne out fully by her behavior. The test comes during the long
bivouac-change march in the wild when the queen must make her
way over a route generally much longer than 100 meters, passing
along a narrow and tortuous trail full of hazards such as vine
bridges and sharp turns although she is swarmed under at nearly
every step by a frenzied crowd of workers.

On a number of occasions the hamatum queen’s participation
in the colony movement has been observed, always at night except
in two instances when colonies were greatly oversize and the move-
ments unduly extended. The bivouac-change usually gets under
way before dusk and is completed during the night, with the
queen coming along during the latter part and frequently near the
very end of the trek. The explanation seems to be that she is
normally stationed in the innermost recesses of the cluster, cen-
tered in the brood with a mass of workers minor around her, in a
section of the bivouac which is likely to be almost the last to be
drawn into the evacuation.

Just before the queen appears from the bivouac there is a very
noticeable increase in excitement among the workers on the trail,
agitated variable movement becomes common instead of the
monotonous plodding seen previously, and the column begins to
widen perceptibly from the 2-3 cm. which is characteristic. As

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the column becomes more crowded it broadens to as much as 15
cm. within a few minutes, then the queen appears in the thickest
part. As the queen moves along she is not only surrounded by
jostling workers of all sizes, but much of the time is literally cov-
ered with them, has them underfoot and hanging to her. The
workers hamper her movements particularly when she slows in
mounting steep grades or in rounding sharp corners ; at such times
they may crowd around so that she is actually stopped for a time.
The remarkable fact is that amid this great excitement and
arduous labor the queen is able to follow the trail in its devious
windings and through its difficult sections.7 Tests show that this
ability depends upon a response to Eciton chemical. For exam-
ple, if the queen is taken up and returned to the route when there
are no workers upon it, she follows it readily. There can be little
doubt about the fact that the queens of Eciton s. str. participate
quite actively in the bivouac-change movements of their colonies
and get to the new site under their own power.

The adaptive significance of the workers’ behavior toward the
queen during the lengthy migratory trek deserves mention. Since
at all times a high pitch of excitement is maintained in the queen ’s
section of the march, any intrusion is immediately responded to
by large numbers of frenzied workers so aroused that they bite
and sting with very little provocation. In the darkness of night,
the feverish activity that prevails in the few meters of column
near the queen contrasts strongly with the lethargic monotonous
lock-step movement that prevails throughout the other sections of
the movement. Only in the entourage of the queen are the work-
ers aroused to a degree of excitement and responsiveness that
resembles their daytime activity in raiding. The outcome is that
the colony’s reproductive specialist is well protected at the time
when exposure is great.

The strength and vitality of the queen are shown impressively
when the colony is etherized for capture. Because of the agility
and speed with which she is capable of moving when disturbed,
the queen generally is able to make her escape from the bivouac
while the principal mass of workers succumbs to the anaesthetic.

7 Reichensperger (1934) observed similar occurrences in a bivouac-change
movement of an E. lucanoides colony in Costa Rica from which he captured
the queen.

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Thus she is frequently discovered under a leaf or piece of bark
near the outer edge of the sheet used to imprison the bivouac, still
somewhat active although workers in numbers lie prone around
her, and if the way is not blocked she may make good her escape
along a raiding trail.

Wheeler (1921) has remarked upon the possibility that an ex-
tensive development in the respiratory system of the Eciton queen
is an important adaptation to a sequestered bivouac life. The
pronounced development of this system in general is indicated by
the prominence of the spiracles, the external openings of the
breathing tubes. With a highly efficient mechanism for gaseous
exchange, it is possible for the queen, despite her huge bulk and
relatively great oxygen needs, to remain for long times within the
close air of the bivouac without any apparent detriment to health.
Furthermore, the large size of the main tracheal vessels of the
abdomen and the extensive ramifications of these vessels among
the reproductive and other visceral organs indicates that the queen
is well equipped in this respect to meet the crucial demands of a
greatly increased metabolism during her brief gestational periods.

Although nothing is known concerning the Eciton queen’s nor-
mal span of life with her colony, in all probability it is equal at
least to one rainy season. However, the hamatum queen for some
reason is particularly short-lived after removal from the midst
of her colony. Three of our 13 contracted specimens lived only
two days and only one lived longer than seven days in captivity
after capture (see Table I). As a control, one queen (1938 B)
was permitted to remain with her colony for more than two
weeks in the laboratory, then was removed (without ansesthesis)
in good condition, yet this queen died within four days after she
was segregated with a small group of workers. Removal from the
colony thus seems to introduce some change which makes in-
evitable the early death of captive Eciton queens. Although in
this study various preliminary attempts were made to discover
the nature of the lethal factor, the results were not sufficiently
clear to warrant discussion here.8 Whatever this factor may be,

8 A possibility worth testing is that the Eciton queen when isolated from
the mass of her colony is deprived of some essential food substance. For
instance, Zahl (1939) has found that workers and queens of the tropical
ponerine ant Dinoponera grandis sicken and die after a few weeks in cap-
tivity unless a larval brood is present in the colony.

June, 1944]

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163

it appears to be specific for queens and not for workers, since in all
cases workers remained alive in captivity for weeks after their
queens had died.

From the above description of workers’ responses to the queen
during the bivouac-change movement, it is apparent that she must
exert a powerfully attractive stimulative effect upon them. The
basis appears to be mainly chemical. Even to the human observer
the queen is distinguished by a delicate, fragrant odor, quite un-
like the heavy, somewhat foetid odor characteristic of Eciton
workers. Workers are especially attracted to pieces of cardboard
formerly in use as floors for the queen’s cell, and will collect in
the one of two compartments in which a queen has rested for a
time. In artificial nests they gather about her, and follow closely
as she runs about. When she comes to rest they pile the brood
around her, and spend much time licking her and stroking her
with antennae. Our queens, placed in small wire cells at the top
of the large cylindrical nests in which their respective colonies
were clustered, were visited by large numbers of minim workers
which penetrated the fine screening and remained to cluster in the
cell. Thus the queen may be considered a factor of some impor-
tance in normal trophallaxis (Wheeler, 1928), the stimulative
interrelationship of individuals which provides the basis for social
organization.9

Characteristics and behavior of the physogastric queen. — At
the time this study began a number of queens of Eciton s. str.
species had been captured in various parts of Central and South
America: notably E. burchelli by Wheeler (1921) in British
Guiana, E. vagans by Gaige in Colombia (Wheeler, 1921), E.
hamatum by Wheeler (1925), and E. lucanoides and E. mattogros-

9 However, it is doubtful that the workers in general are widely sensitive
to the actual presence or absence of the queen as Wheeler (1921) implies they
were in the case of a colony of E. burclielli from which the queen was re-
moved. “I infer,” he says, “that this was the only remaining female in
the colony, for after her removal a perceptible apathy or dejection seemed
to fall on the whole body of ants” (p. 298). Actually, this effect of
1 1 apathy, ’ ’ specifically a rather abrupt drop in general activity, characteristi-
cally ensues after large numbers of the ants have been stirred up and then
permitted to recluster. Once reclustered, they fall into a lethargic condition
whether or not the queen has been removed, hence this behavior cannot be
attributed to a removal of the queen.

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sense by Reichensperger (1926, 1934) and E. quadriglume and
E. rogeri in the Argentine by correspondents of Carlos Bruch
(1934). Thus by 1934 the queens of nearly all of the species of
Eciton sensu stricto had been taken. It is interesting to note that
in all of these cases the queen’s gaster was contracted, a circum-
stance which led various writers ( e.g Bruch, 1934) to conclude
that the queen in question was “young,” or “virgin.”

The experience of the present writer was similar in capturing
Eciton s. sir. queens in Panama for behavior studies. In 1933 two
queens, E. hamatum and E. burchelli, were taken; in 1936 four
queens, one of E. lucanoides and three of E. hamatum, and in
1938 nine more of the last species, all of them alike in the fully
contracted condition (see Table I) . Prom these and other facts it
becomes apparent that few if any of the queens of Eciton s. str.
reported in the literature could have been virgin when captured,10
and that the dichthadiigyne must be capable of returning to the
contracted state after having delivered eggs. This interpretation
in fact was offered as one alternative by Wheeler (1925) in report-
ing the first capture of the queen of E. hamatum.

Early in the present investigation, evidence concerning periodic
changes in colony behavior, together with the experience of cap-
turing numerous queens singly from colonies with broods in all
stages of development, strongly indicated that the queen must
pass repeatedly from the contracted to the physogastric condition,
remaining in the latter condition only briefly before returning to
the contracted state. Finally it became clear circumstantially
that the eggs of a given brood must be laid during a short period
of a few days near the end of the first week of the statary phase
in colony behavior, when the colony is non-migratory.

Following this conclusion, in 1938 an effort was made to capture
a hamatum colony at the critical time, which on circumstantial
grounds was placed about seven days after the beginning of the
statary period. On July 21 a colony (record number 38 H)
was found which although nomadic at the time was evidently
nearing the statary phase, judging from the fact that its larval
i° Probable exceptions are the two queens of E. burchelli taken by Wheeler
and Emerson at Kartabo, British Guiana, in 1920 (Wheeler, 1921), in which
circumstances point to the recent emergence of at least one and perhaps both
of them from cocoons.

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165

brood seemed very advanced and about ready to spin cocoons.
After three further bivouac-change movements on successive
days, when spinning appeared to be mainly finished in the brood,
the colony settled down at a spot near station 4, Barbour trail.11

The statary bivouac was formed in a deep cranny near one end
of a decayed fallen tree-trunk. The cluster formed a long ellip-
tical curtain-mass filling the opening of the niche. On each of the
following days, when the colony was revisited, the ants had only
one raiding-system, a typical sign of the statary condition. The
site was revisited on July 30, i.e., one week after the beginning of
the statary period, with materials for capturing the colony.

The colony had withdrawn about 30 cm. farther into the recess,
very probably in response to frequent spattering by rain. Fortu-
nately it was still fairly accessible. The bivouac was quiet, its
forward wall sprinkled with ants holding cocoons in their mandi-
bles. A dampened sheet was fastened securely across the open-
ing to entrap the ants, then was sprinkled with ether. After
four minutes, when the covering was removed, the major portion
of the colony, a mingled heap of ants and cocoons on the floor
of the cavity, was quickly scooped into a large jar. A few strands
of ants still hanging, together with masses of eggs and clusters of
small workers from the rear of the bivouac, were put into a second
jar. In examining the material promptly after returning to the
laboratory the first jar was found to contain the major part of the
large enclosed pupal brood as well as most of the worker popula-
tion. In the second jar, which contained workers and egg masses
from the rear strands of the bivouac, a phy so gastric queen (Fig.
1) presently was brought to light in the part of the mass where
workers minor and eggs were most numerous. Judging by rela-
tive positions in the jar, this queen must have occupied a position
near the back of the cluster, probably among the hanging strands
which contained most of the workers minor and most of the eggs.
A careful inspection of the material revealed no other queens, no
males, and two broods — a large enclosed pupal brood and an enor-
mous clutch of recently delivered eggs.

Within the following twenty-four hours, which proved to be the

11 The queen was seen in the procession during the final bivouac-change
movement, and was judged to be in the contracted condition at that time.

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span of life remaining to the 38 H dichthadiigyne, her behavior
was studied as continuously as other duties permitted. —

A few minutes after she was removed from the jar (4: 30 p.m., one hour
after capture) the queen appeared to be recovering from ansesthesis, as indi-
cated by reflex twitchings of tarsi and antennas, although most of the workers
remained immobile. At 4 : 45 p.m. she had recovered sufficiently to gain an
upright posture and had laid 20 eggs. At 5 : 10 p.m. she was running about
the Petri dish in which she had been placed, despite the trammeling effects
of her huge distended gaster which tilted to one side or the other as she
moved. From rough measurements at that time, her gaster was 16.5 mm. in
length.

Until 8 : 15 p.m. the queen remained undisturbed in a darkened Petri dish,,
laying more than 300 eggs. Then for an hour she was exposed at intervals
to brilliant photoflood light when motion pictures were taken. Despite evi-
dent disturbance from the light, marked by shock reactions when light was
introduced and by turning from the source, there was a describable regularity
about the queen ’s behavior during this interval. There were successive
periods of a few minutes each in which the queen stood in place, observably
engaged in egg-laying, with intermittent periods in which she ran about the
dish more or less continuously, dragging or carrying numerous workers on
her gaster and followed closely by others. At no time was she moved by the
workers, although generally a number of them remained close to her whether
or not she was in motion.

The following representative notes were taken during two of the quiescent
intervals, the first lasting 5 minutes after 9: 37 p.m. and the second 2 min-
utes after 9: 44 p.m.:

The queen abruptly breaks pace and suddenly stops in place, headed away
from the photoflood source. She stands firmly on all six legs, with the
ventral surface of the gaster resting on the floor. In oviposition, the eggs
emerge in a wide jet from the vaginal orifice, and spread out radially upon
the hypogynium. The process is marked by slight laterad oscillations of the
the gaster and a shivering of the body and legs, which cease when eggs no
longer emerge. — Eggs were laid during both of these halts.

At 11 : 30 p.m. the queen with 6 workers was placed in dim light for obser-
vation. During the next 90 minutes there were 11 quiescent periods, varying
between 30 sec. and 11 min. in duration (6 of them lasting between 1 and 4
min.). The intervening periods of activity were shorter, most of them
around 3 min. in duration. In each case, locomotion began rather abruptly.
The quiescent periods likewise began abruptly, with the queen usually sprawl-
ing close to the floor at once. Generally she stopped in a position facing
away from the light, which although dim was sufficient to orient her. In
some of the shorter intervals no eggs were laid. In the course of one or two
of the longest stops, the queen shifted position briefly or moved forward a
short distance once during the interval. The workers remained close to her,,
frequently applying their mouth parts to her body at the vaginal orifice and

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• Schneirla: Insect Behavior

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at the surface of the hypogynium and licking these surfaces. After one of
the stops a drop of viscous greenish-yellow fluid remained where the tip of
the queen’s gaster had rested. The workers soon found this drop and applied
their mouth parts to it, evidently feeding. — Observations were discontinued
at 1: 30 a.m., since no eggs had been laid during the preceding 30 minutes. —
Between 5: 10 p.m. and 12: 00 a.m., 1245 eggs were laid.

In the morning, at 6: 30 a.m., only 358 eggs were found to have been laid
during the night ( i.e ., after 1: 30 A.M.). As before, alternating periods of
quiescence and of activity were observed. Between 8: 15 and 9: 54 a.m. the
queen was shielded by a ruby-glass filter and remained under observation.
During this interval there were sixteen quiescent periods alternating with
intervals of continuous locomotion, virtually all of the phases of quiescence
and of activity falling between 1 and 4 min. in duration. In some of the
stops between 20 and 30 eggs were laid, in others no eggs. The following
notes typify egg-laying behavior:

8 : 23 to 8 : 26 a.m. — The queen stops abruptly, facing away from the weak
light. Promptly there begins a shivering movement of the gaster, increasing
at times to an oscillatory movement ; then the oscillation becomes continuous,
at times reaching 1 mm. in amplitude. These oscillations of the gaster begin
locally and increase in amplitude, meanwhile spreading to include the entire
body with the legs participating. Near the height of such spasms the eggs
begin to appear. A jet of eggs between 12 and 15 units wide is forced slowly
from the vagina directly backward over the hypogynium, with the eggs
fanning radially to each side as they emerge. The eggs in small packets are
either picked off by the workers or drop to the floor when the queen moves
away/ — Finally the queen breaks abruptly into motion, in an interval of loco-
motion which lasts 4 minutes in this case.

At 4 : 15 p.m. the queen seemed less energetic than in the morning, and had
laid only about 60 eggs since 10: 00 a.m. For about 20 minutes she was
exposed to bright sunlight while photographs were taken, and although she
was shaded between exposures the intense light appeared to exert a decidedly
injurious effect upon her. At length she fell upon one side with flexed legs,
unable to regain an upright position. At 6 : 00 p.m. there were unmistakable
signs of approaching death, which in the contracted queen is characterized
by intermittent struggling and by local tarsal and antennal reflexes lasting
a number of hours. — At 6 : 30 p.m. she was chloroformed for fixation in
Carnoy’s solution and eventual preservation in 70% alcohol.

Notwithstanding the fact that disturbances incident to labora-
tory captivity must have disrupted the normal oviposition
mechanism greatly, and although the factors which character-
istically hasten death in captive Eciton queens must have been at
work, certain inferences concerning the normal circumstances of
physogastric function may be drawn from these observations.
Concerning the queen herself, the regular occurrence of short

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intervals of action and of quiescence (and egg-laying) in alterna-
tion indicates that a rhythmic process underlies the delivery of
eggs from the ovarioles. The abrupt onset of each quiescent
period together with the oscillatory movements of the gaster and
at times the entire body during oviposition speak for the opera-
tion of a fairly distinct egg-laying process arising at short inter-
vals. Likewise, the abruptness with which the queen broke into
motion after each quiescent period indicates the regular recur-
rence of a rhythmic change in this process, evidently a phase dur-
ing which more eggs became viscerally available for delivery.

Under the conditions of these observations the queen was at-
tended by only a few workers, and space permitted her to run
about in the intervals between egg-laying stops. It is probable
that under normal conditions in the bivouac, when she is sur-
rounded by masses of workers, free locomotion is out of the ques-
tion for the queen. In that situation her viscerally-stimulated
overt activity must be reduced to stirring in place and to inter-
stimulative relations with workers.

This queen died with a considerable portion of her eggs still
undelivered, a fact which together with other circumstances indi-
cates that in all probability lethal processes setting in with cap-
tivity impaired the egg-production rhythm seriously from the
beginning. Under normal conditions in the bivouac this rhythm
presumably functions smoothly and continuously throughout the
period of approximately four days in which all of a given clutch
of eggs is delivered. Then, as we shall see, circumstances are such
that the queen, contracted once more but in an exhausted condi-
tion, may profit by a long interval of recuperation before the
parturitive ordeal must be repeated.

Our results shed some light upon the behavioral relation of the
physogastric queen to her colony. First of all, the fact that the
queen is quite capable of locomotion during her time of physo-
gastry effectively negates the hypothesis (Muller, 1886) that the
colony must remain in place at such times because of an inability
to drag along the heavy egg-laden queen. She moves readily, and
even drags or carries workers about with her, hence there is reason
to believe that she would be quite capable of joining a bivouac-
change movement if one happened to get under way. She would

June, 1944]

Schneirla: Insect Behavior

169

have to make periodic (viscerally enforced) stops on the trail,
but could make the trek even though it might well be fatal. We
shall find that the sessile (i.e., statary) status of the colony when
the queen is laying her eggs is indeed related to the queen’s
oviposition cycle, but very indirectly, and very differently than
the above hypothesis would suggest.

Our observations indicate that the normal stimulative attrac-
tiveness of the queen for the workers (i.e., her trophallactic rela-
tionship with workers) is considerably increased during the time
of oviposition. The intersegmental membranes of her enormously
swollen gaster and the hypogynium and vaginal regions in par-
ticular are highly effective centers of attraction. The workers are
constantly crawling upon her and licking these regions as well as
the eggs when they appear (cf. Wheeler, 1900) and the (evidently
much relished) small drops of fecal material which issue at times
after egg-depositing episodes.12

There can be little question that this queen produced all of the
eggs taken with the colony, a total of 17,062, which was estimated
to be two or three thousand short of the actual number laid prior
to capture. To this we may add 2,046 eggs laid while the queen
was held captive, and 7,190 unlaid eggs taken from her gaster in
post-mortem study. The total of 26,298 eggs approaches fairly
close to the size of Eciton brood populations for which census
studies have been made in other cases, offering further evidence
that one individual dichthadiigyne is capable of delivering single
broods of that size. In view of circumstances indicating that the
eggs of each huge brood are laid within a short interval of prob-
ably no more than three or four days, the entire performance
seems quite worthy of being termed a stupendous feat.

The great susceptibility to death of the Eciton queen in this
condition is attested by the fact that our specimen survived only
about 30 hours after capture. It is probable of course that her
demise was hastened by the anassthesis and by other shock-effects
( e.g ., from intense light during photography), yet the fact that
contracted queens similarly treated usually remain alive for as
long as a week in captivity speaks for a specific physiological
vulnerability to non-optimal conditions in the gravid Eciton

12 Emerson (1939) has reported a similar observation for queen-worker
relationships during egg-laying in termites.

170

Journal New York Entomological Society

t Vol. LII

queen. It is a fact of great adaptive significance that during this
biologically critical time the dichthadiigyne ordinarily is assured
the essential optimal situation through the operation of intrinsic
factors which render the colony sessile. No less remarkable is the
fact that the queen herself , through her peculiar reproductive
properties , is indirectly the basic controller of this state, of affairs.

EVIDENCE FOR A FUNCTIONAL RELATIONSHIP BETWEEN
BROOD AND COLONY

Significant periodic differences in colony behavior. — As the
first step toward working out relationships between the queen and
her colony, let us examine the general situation of the series of E.
hamatum colonies from which queens were captured for this study
(see Table I).

In their behavior twelve of the hamatum colonies (i.e., colonies
33 A; 36 A, B, D, and G; and 38 C, X, D , E, F, G, and I) from
which queens were taken conform to the pattern previously de-
scribed as “ nomadic’ 9 (Schneirla, 1938). On the other hand
only three (i.e., colonies 38 A, B, and H) conform to the pattern
described as “statary. ” Since all of these colonies were studied
for at least three days prior to capture, and some of them for
longer times, any important deviations should have become ap-
parent ; but in all cases there appeared very clearly the character-
istics of one or the other of the described activity patterns.

In cases showing the nomadic pattern, all of the colonies were
migratory, that is, they had all engaged in bivouac-change move-
ments near the end of each day in the period preceding capture.
Without exception among many observations, the raiding activi-
ties of a given day were terminated by a complete movement of
the colony, generally during the evening and early night, so that
the next day found the colony developing a new raid from a dif-
ferent site generally more than 100 meters removed from the last.

Although their bivouacs were situated in rather different topo-
graphical circumstances, the clusters of the “nomadic” colonies
were not secluded as is typical of statary colonies. The former
almost without exception established themselves beneath logs,
under matted vines, or against the sides of trees between but-
tressed roots, seldom within cavities such as hollow logs or trees.
As a rule the larger portion of the more or less cylindrical mass

Data Relevant to the 15 Queens of E. hamatum Captured in the Present Study, and to the Colonies from

Which These Queens Were Taken ( cf . Fig. 2)

June, 1944]

Schneirla: Insect Behavior

171

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* The growth stage of the respective broods taken in 1938 is represented by data on the range (E) and the mean (M) of
body lengths (samples ranged between 25 and 200 specimens from each colony). The writer wishes to express his thanks
to Miss Ruth Greene, who generously contributed her time to gather these data.

The larvae of various Eciton species have been described by G. C. Wheeler (1943) with a key for their identification.

172

Journal New York Entomological Society

[Vol. LIT

of ants was fairly exposed to view. It is typical that the bivouacs
of nomadic colonies are far more readily approached and observed
than are those of statary colonies.

In their raiding, particularly, colonies in the two behavior con-
ditions exhibited very different characteristics. In the nomadic
condition the Ecitons always staged vigorous daily raids which
began promptly with the first daylight and grew rapidly into the
characteristic maximal raiding pattern of the species. In E.
hamatum this is marked by the growth of two or three (generally
three) principal raiding systems, each a tree-like pattern of
trails with a single principal trail as its line of communication
with the bivouac. Although of course the details of the trail
systems vary considerably according to the situation of the
bivouac and the general topography, invariably multiple trail-
systems are found under nomadic conditions. At such times a
maximal number of ants from the population is drawn into the
daily foray, developing extensive and complexly branched trails
on a widespread front of raiding. With the raid probing out
along a number of principal lines, the usual result is that roughly
three-fourths of the circular zone around the bivouac is invaded,
with the consequence that relatively enormous quantities of booty
(mainly the soft-bodied young of other insects) are gathered in.
While nomadic , a colony remains near its peak of vigor in raiding
and in other activities.

With those colonies in the statary condition when their queens
were captured the case was quite different in all important
respects concerning behavior. Colonies 38 A, B, and H were
all known to have remained in their same bivouacs at least three
days prior to capture. Colony H, for example, after three ob-
served daily bivouac-changes, clustered within a large crevice
in the side of a massive log on July 22, 1938, and was found in
the same place and in almost the same spot on July 30, when
captured.

Other evidence has been offered (Schneirla, 1933, 1940) for
the existence of the statary period as a distinctive 4 1 sessile ’ ’ phase
of Eciton life. Observations on particular colonies extending
over considerable periods have shown that once a hamatum colony
enters a statary phase, it remains bivouacked in the same place

June, 1944]

Schneirla: Insect Behavior

173

and exhibits other characteristic “statary” features of behavior
during a period of approximately 19 days, before a major change
occurs.

The fact will be recalled that of the fourteen colonies from
which queens were captured in this study, only three were taken
in the statary condition. This difference is scarcely a matter
of accident, but is due to the relatively secluded locations taken
by colonies of Eciton s. str. when they are statary. Colony 38 H,
clustered in a deep crevice on the outer wall of a huge log, was
unusually accessible for a statary colony. The virtual impregna-
bility of statary colonies is attributable to two facts in particular.
First, they are commonly clustered within a cavity, generally
in a hollow log or tree. Colony 38 A had to be extracted from a
hollow log, colony 38 B from within the large hollow root of a
standing tree, by drilling a circle of holes with brace and bit
and breaking out an opening through which the ants could be
removed by hand.13 Then, too, if a statary colony happens to
settle where it is somewhat exposed to the elements, disturbance
from wind, rain or sunlight causes the ants to shift their position
so that after a few days capturing them would require the assis-
tance of a wrecking crew. The proneness of colonies to cluster
in hollow logs or trees when entering the statary period stands
as a highly adaptive circumstance for which no specific explana-
tion can be advanced at the present time.

Without exception, each of our three statary colonies from
which queens were obtained had a single raiding system when
captured. This pattern is readily recognized by the presence of
just one principal raiding trail leading from the bivouac, rather
than two or three as in the nomadic phase. Distance reached
from the bivouac in the raiding is not a secure criterion of its
relative vigor, since in the statary phase the chemically-marked
route of a previously used trail may be employed on more than
one day so that such trails often become greatly extended beyond
the distances ordinarily reached in nomadic-phase raiding. As
more certain indications of reduced raiding, the number of prin-
cipal systems is one rather than three, and there is a smaller

13 Dr. Neal Weber kindly assisted in this operation, which was completed
in a torrential rain.

174

Journal New York Entomological Society

[Yol. lit

number of branch trails in that system than is characteristic of
' a raiding system in the nomadic phase. Thus, in the statary
phase, raiding activity falls much below the level of vigor and of
numbers involved during the nomadic phase.

Before considering the relationship of these periodic differences
in Eciton behavior, it is desirable to examine correlated differ-
ences in the internal constitution of the colony.

Intrinsic conditions underlying colony behavior. — A considera-
tion of the internal characteristics of the colonies (see Table I)
shows that the external behavior differences were paralleled by
others no less outstanding. The intrinsic differences may be
represented by a comparison of the two groups of colonies in
terms of their condition and the status of their brood or broods
when the queens were captured.

First of all, it should be emphasized that the broods invariably
were worker broods, and that no batch of larvae or pupae con-
tained individuals departing sufficiently from the others in form
or size to suggest the anlagen of fertile forms. This statement
is of course not based upon our present facts alone, but finds its
main support in an examination of hundreds of Eciton broods
on Barro Colorado Island in rainy months between May 15 and
September 20 in four different years which has disclosed none
but worker forms in them.

Another fact of significance is that while each of the broods
contained tens of thousands of individuals, all members of a given
brood were roughly at the same stage of development. This con-
dition held not only when a single brood in the larval stage was
present, but also for cases in which there were two broods (i.e.,
a pupal brood in addition to a brood in the very early larval
condition). This statement, likewise, is based not only upon the
present data but also upon the invariable finding that in E.
hamatum the entire population of a given mature larval brood
terminates that stage and spins cocoons within a very few days,
and that pupal broods mature and are removed from their cocoons
as callows within a similarly short period of three or four days.

All three of our statary colonies had two broods at well-sepa-
rated points of development, a fact which we shall find character-
istic of the latter part of this activity phase. Colony 38 H con-

June, 1944]

Schneirla: Insect Behavior

175

tained a well-advanced pupal brood as well as a massive batch
of newly laid eggs. Colony 38 A contained a still more advanced
pupal brood, and its younger brood had largely hatched and
passed into the early larval period. The two broods of colony
38 B corresponded in general to those of 38 A. In the early days
of the statary period only one brood (always a pupal brood)
is present.

In contrast, the nomadic colonies each contained only a single
brood, in every case in the larval condition. From the size data
on these broods, represented in Table I by the averages and ranges
of individual body lengths, it is apparent that the broods were
at very different stages of development when the colonies were
captured. If a greater magnitude of body length in the brood
(as indicated by larger averages and greater ranges) may be
taken as valid evidence for an advanced point in growth, the
nomadic colonies captured in 1938 may be placed in the following
sequence : X , C, D , E, F and G. It will be noticed that with the
single exception of colony X, there is a correspondence between
the chronological order of capture and the growth point attained
by the respective broods. The possible significance of this fact
will be considered in a later connection.

When we bring together our facts concerning the external and
internal characteristics of the colony, a significant parallelism is
evident between them for both the nomadic and the statary
phases. In the statary phase 1) the colony does not migrate
but remains in a given place, 2) by virtue of its secluded location
it is well protected from the elements and from animals, 3) it
raids minimally and is comparatively lethargic, and 4) it con-
tains a pupal brood and in its latter part ( e.g ., colonies 38 A,
B, and H) eggs or a young larval brood as well. The most sig-
nificant parallel circumstances are a low condition of colony
activity and a brood {or broods) incapable of overt activity.

In sharp contrast to the above situation, a colony in the no-
madic phase 1) regularly moves to a new bivouacking site at
the end of each raiding day, 2) generally bivouacs in fairly open
places, 3) raids maximally and appears to be always {i.e., in
the daytime) near the peak of activity, and 4) contains a single
brood in a more or less advanced larval phase. In this case, the

176

Journal New York Entomological Society

[Vol. LII

most significant parallel circumstances are a high point in colony
activity and the presence of a larval brood which is capable of
overt activity .

Theoretical explanation of the parallelism between brood con-
dition and colony behavior. — These facts suggest that a causal

DAYS

Fig. 2. Schema to represent concurrent events in colony behavior (top),
condition of the brood, and function of the queen in E. hamatum. (The
sequence of given events in the colony may be traced horizontally, the con-
currence of events, vertically.) A-H, approximate situation of the respec-
tive colonies from which queens were captured in 1938 ; Ph, physogastric
interval of the queen; E, egg mass delivered in given physogastric interval;
L, larval stage of brood; P, pupal stage of brood; £ 2J, workers minor and
major of a callow brood.

relationship exists between the condition of the brood and the
circumstances of general activity in the colony. The conclusion
appears justified that conditions arising from the capacity of the

June, 1944]

Schneirla: Insect Behavior

177

brood for overt activity somehow account for the critical differ-
ence which appears in general colony activity pattern when active
broods rather than “passive” broods are present. This idea
has been advanced in a previous connection (Schneirla, 1938)
and will be carried further presently.

Evidence from field observation and special test supports this
inference of a causal relationship dependent upon condition of
the brood. Batches of eggs and very young larval broods (as in
the statary colonies 38 A, B, and H) are as a rule gathered itito
a single mass or a few masses which are covered and at times per-
meated by minim workers and are only very indirectly in contact
with the adult population of the colony. On the other hand, when
the larvae have developed further (as in colonies 38 C, D, E, F,
G, and X) and their twisting movements become perceptible to
the unaided eye, these masses are broken up and more or less dis-
tributed through the bivouac, so that larval activity and other
stimulative effects may directly reach large numbers of adult
workers. When the larvae mature, and become enclosed in co-
coons, it is apparent that in the ensuing pupal stage (as in col-
onies 38 A, B, and H) they become passive contents of the bivouac.
Heaped in interior pockets of the cluster and held by workers
distributed through its wall, they are presumably neutral in im-
portance or may actually exert a quieting effect upon the work-
ers that clutch them in their mandibles, somewhat as the pressure
of thumb-in-mouth pacifies an infant.

From laboratory observations we may say that through the
presence of an active larval brood the workers are distinctly
stimulated so that the amount and the vigor of their activity in-
creases greatly. When Ecitons are divided into part-colonies of
comparable size, it is the groups with larvae that show the greater
activity both within the confines of their artificial nest and in the
frequency of visits to an adjoining food-place, as compared with
groups that have no brood or have a pupal brood in cocoons.
When we study the relationships of individuals within the nest,
there are clear indications that this difference in activity level
actually depends upon a stimulative effect from the larval brood.
Typically the larval brood is spread over an area, usually around
the queen if one is present. The workers move restlessly around,

178

Journal New York Entomological Society

[Vol. LII

frequently touching the larvge with mouth-parts or licking them,
intermittently touching and stroking them with antennae, often
picking them up and shifting their position slightly or carrying
them bodily from place to place. When pieces of food are car-
ried into the innermost nest area, they are commonly deposited
near the larvae or actually laid upon them, so that in their almost
incessant twisting movements the larvae readily bring their mouth
parts into contact with the food.14 It is a common observation
that stimulation from passing workers may arouse quiescent larvae
to squirming activity, or that chance stimulation from larvae may
bestir workers. It is manifest that the larvae thus in a number
of ways arouse the adults to increased activity, in the mutual
stimulative relationship between adults and brood which Wheeler
(1928) has termed ‘ ‘ trophallaxis. ”15
On the other hand a group without any brood or with pupae in
cocoons is distinctly less lively than when larvae are present.
Even when exposed to light the workers seldom appear very ex-
cited, and most of the time they stand quietly over their heap
of cocoons or near it, or move slowly about in the vicinity. Under
these conditions Ecitons are chronically less feverish in their ac-
tivity, indulge in many fewer excursions from the nest, and hence
they carry in far less food than in the larva-containing situation.
We may say that the group or colony which contains active larva
as in the nomadic period has a special source of colony “drive.”
In contrast, in the statary period, this factor is greatly reduced
when interindividual stimulation is limited to relationships among
adult workers themselves.16

14 Gallardo (1920) has commented upon the great mobility of Eciton
larvae.

is The gastric anatomy of the larvae of B. burchelli and the feeding be-
havior of these larvae have been described by Wheeler and Bailey (1920).
In this species the larva is fed at frequent intervals on boluses composed of
the soft parts of insect prey, laid by the workers upon the ventrum of the
larva or actually thrust into the larval gullet.

i6 From his observations on a colony of B. burchelli, Wm. Muller (1886)
inferred that an increase or decrease in the raiding activities of a colony is
attributable to the relative need of the brood for food.

“Larven brauchen ja im allgemeinen, besonders kurze Zeit vor
der Yerpuppung, bedeutend mehr Nahrung als die fertigen Insekten,
und so scheint nichts natiirlicher, als dass das Nahrungsbediirfnis der

June, 1944]

Schneirla: Insect Behavior

179

However, in a statary colony with advanced pupae there is a
marked change in the general level of activity when the nearly
mature individuals begin to move within their cocoons. When a
close approach to the end-poinjt of pupal development is indicated
by the appearance of a distinct pigmentation of eyes, mandibles,
and anterior sections of the body, activities such as twitchings of
the legs and slight turnings of the body become observable. Such
movements of the pupae may be increased by picking individuals
up with tweezers or by lightly probing the body, just as they are
perceptibly elicited when the cocoons are picked up by workers.
The stirring of mature pupae appears to excite the workers and
evidently leads to the removal of the brood from cocoons ; the en-
velopes readily tearing open as the workers manhandle them
(Schneirla, 1934, pp. 320-321). That the opening of cocoons de-
pends critically upon pupal movements is suggested by the fact
that test groups did not open cocoons in which the advanced pupae
had been killed with a needle (op. cit., 1938, p. 66).

There is a distinct increase of excitement in an Eciton colony
as emergence of its brood gets under way. The excitement rises
in crescendo until, a day or two later when the largest part of the
brood is free and active as callow individuals, the colony shakes
itself from the statary period by staging an exceptionally vigor-
ous raid ending in a bivouac-change movement. Laboratory ob-
servations show that in the appearance of a new pupal brood and
in the strong stimulative effects from callows during their first
few days as “free” individuals in the colony, an Eciton worker
population receives a great lift in internal “drive.” The conclu-
sion seems in order that this internal change is responsible for
setting a colony into the nomadic phase.

Gesellschaft ein geringeres wird und die Tiere entsprechend weniger

auf Beute ausgehen, nachdem alle Larven eingesponnen 7 ’ (p. 87).

While this point undoubtedly has relevance to the 'phylogenetic problem (con-
cerning the evolution of the Eciton behavior pattern) and describes an
important adaptive function of the pattern, it is teleological and misleading
if applied to the ontogenetic problem (concerning the contemporary causa-
tion of the pattern in a given colony). As the present discussion shows, it is
not the brood ’s food-consuming capacity as such, but the presence or absence
of incidental tactuo -chemical stimulation from the brood which critically
governs periodic changes in colony behavior.

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Journal New York Entomological Society

[Vol. Lll

Our theory thus accounts for the nomadic-statary (rainy-sea-
son) cycle of Eciton colony activity in terms of cyclical changes
in the internal stimulative properties of different broods. In
Figure 2 the relationship is represented schematically in terms
of a diphasic diagram, on which are indicated the positions evi-
dently attained by our colonies at the times their respective
queens and broods were captured. The “trough” phase of the
cycle signifies the statary period, in which the colony is held in
position and is comparatively inactive and the brood or broods
relatively passive in their social roles. A colony enters this
statary phase when the enclosure of its mature larval brood re-
moves the major internal source of special social stimulation.
The appearance of a new brood as eggs laid after about one-third
of the period has elapsed does not materially change general
colony behavior, since with this brood concentrated in masses,
incapable of movement, it can exert only a relatively negligible
stimulative effect upon the general population. As a matter
of fact, the vigor of raiding seems to decrease somewhat toward
the middle of the statary period, and hamatum colonies sometimes
show raidless days then (also E. burchelli, as Muller (1886)
noticed). With social stimulation low throughout the period, the
workers are only weakly stimulated to leave the colony in raiding,
with the result that statary-period raids are relatively feeble as
compared with nomadic-period raids.

We may say that in the statary period a low summation of
colony excitement directly expresses its limitations in the under-
development of raiding, which roughly is only about one-third as
extensive as in the nomadic period. Such raids may be regarded
as underdeveloped not only in their restricted numbers and scope
but also in falling below the threshold of development at which
a bivouac-change movement becomes inevitable.

To speak of a “threshold” of raiding essential for new behavior
requires us to show why underdeveloped raids cannot lead into
a colony movement. This has been done in special studies of the
relationship between raiding and the bivouac-change movement
(Schneirla, 1938, 1944) which bring out the reason why a colony
can withdraw to its original bivouac from an underdeveloped
statary-period raid but must move onward when a maximal

June, 1944]

Schneirla: Insect Behavior

181

nomadic-period raid has been staged. The actual behavior proc-
ess, fairly complex, centers around the fact that in the statary
period the colony is insufficiently rearoused (by regular post-
meridional atmospheric changes) after the midday lull in ac-
tivity, so that no exodus exists capable of blocking the principal
trail to returning raiders. In the nomadic period, on the other
hand, the afternoon rearousal produces a lasting afternoon exodus
from the bivouac which forcibly reorients returning raiders out-
ward in at least one trail system; then a movement from the
bivouac persists on this route, eventually draining the entire
colony to a new bivouacking site. Thus when extrinsic stimu-
lation (from light in particular) drops away at dusk, a colony in
the statary phase loses its principal source of arousal and grad-
ually approaches quiescence; whereas a colony in the nomadic
phase when raiding declines with dusk possesses an adequate and
persistent internal “ drive’ ’ facilitating the vigorous exodus al-
ready in progress. The key to the situation appears to be the
radical difference in the intrinsic properties of colonies in these
two conditions.

The break from the statary phase appears to be accomplished
not through the influence of the new larval brood, which at that
time seems insufficiently developed to be a social-stimulative
factor of any account, but rather through activities centering
around the maturation of the pupal brood and its appearance as
an enormous horde of hyperactive, voracious callows. Invari-
ably, hamatum colonies shift from their statary sites when the
pupal brood has largely emerged; the deserted spot is littered
with empty cocoons, and the first bivouac-change processions are
distinguished by crowds of pale-colored callow workers. Circum-
stances indicate that the stimulative effect from tens of thou-
sands of callows not only serves to bring the colony into nomadic
life, but also persists for a few (apparently about five) days as
a factor of similar influence and importance. Our colony 38 X
was taken at such a time. As a major source of stimulation, the
presence of the callows appears to overlap the involvement of the
new brood of larvae, in that as the callows lose their early social-
stimulative effect and merge into the adult worker population, the
larvae become increasingly functional in this respect. About five

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Journal New York Entomological Society

[Vol. LIT

days after the nomadic period has begun, typically, the callows
are almost indistinguishable from regular workers and (judging
from laboratory observations) have dropped to the level of the
ordinary workers in their trophallactic properties. Meanwhile,
the larvae have increased notably in size and activity, most of them
are capable of feeding, and at this time they appear to constitute
the new major source of social stimulation or intrinsic colony
‘ ‘ drive. ’ ’ Our colony 38 C was taken at such a time. When it
was captured, the larval brood was distributed rather widely
through the interior of the bivouac, and in the laboratory a pro-
nounced activity was noted in all but the smallest size category.

Colonies 38 C, D, E, F, and G (taken in chronological sequence)
may be placed at successive points in the nomadic period on the
basis of the invariable fact that an increasing development of the
larval brood parallels an advance through this phase of activity
(see Fig. 2). Not only are the trophallactic properties of the
brood instrumental in maintaining the nomadic pattern of be-
havior by supporting daily maximal raids, but as the larvae de-
velop further their influence appears to increase. The apparent
augmentation of their social effect probably is due to increased
chemo-stimulative properties as well as to greater general activity
as they grow. This is suggested by laboratory observations and
by the fact that in the bivouac more advanced broods tend to be
widely distributed through the cluster, single larvae held by indi-
vidual workers much more frequently than with younger broods.
Thus the intrinsic ‘ ‘ drive ’ ’ effect of the brood not only maintains
the nomadic pattern in the colony, so that maximal raids and
bivouac-change movements appear regularly in the daily routine,
but this relationship appears to rise through a crescendo as brood
development nears its climax. Hence, near the end of the no-
madic period, raids are more extensive and more heavily popu-
lated, and colony movements tend to carry over longer distances
than earlier in the period. In particular, this is revealed by
studies of individual colonies throughout the nomadic period
( e.g colony 1936 A, — Schneirla, 1938; also E. burchelli, colony
381, — Schneirla, 1944).

As we have pointed out, in addition to postulating an excitation
to “maximal’ raiding through the drive effect of active broods,

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Schneirla : Insect Behavior

183

the theory postulates a given threshold in raiding which is essen-
tial for the occurrence of a bivouac-change. As the relationship
has been worked out in a previous paper {op. cit., 1938), only in
the nomadic period does raiding attain the pattern {i.e.y three
trail systems in hamatum) and the degree of colony involvement
that is required if it is to eventuate in a colony movement. That
the essential “maximal” pattern of activity in the colony depends
specifically upon a characteristic larval-brood influence is sug-
gested strikingly by the regularity with which colonies lapse into
the statary period on the very day most of the larvae have spun
their cocoons.

It seems clear that since a critical difference in the pattern of
hamatum colony behavior depends upon the condition of the
brood, the ultimate determination of such matters resides in the
individual that produces the broods, i.e., the queen.

THE ECITON QUEEN AS PACEMAKER FOR COLONY BEHAVIOR

It is characteristic of E. hamatum, E. burchelli, and other rep-
resentatively terrestrial species of Eciton s. str. (if not Ecitons
generally), that broods appearing in the rainy season are very
large, and that all individuals in a given brood develop concur-
rently. That is, in each new hamatum brood the eggs are laid
within a span of a few days, pass through the larval period essen-
tially in step, enter the pupal stage within about four days, and
appear as callow workers all within a very short period. Obvi-
ously this state of affairs depends essentially upon the reproduc-
tive properties of the Eciton queen.

Let us first consider some further facts about brood develop-
ment that throw light indirectly upon the capacities of the queen.
Population studies have been made with four broods captured on
Barro Colorado Island, with these results : E. hamatum — a larval
brood, 26,452 individuals (not complete), a pupal brood, 31,379
individuals (fairly complete) ; E. burchelli — one complete larval
brood of 36,888 individuals, and a pupal brood of 31,298 indi-
viduals. Then too, the mass of eggs from physogastric queen
38 H totalled more than 26,000, probably with a few thousand
missing. Since many other broods inspected in the egg, larval, or
pupal condition were comparable in bulk with these populations

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which were actually counted and sized, it seems that the rainy-
season broods of E. hamatum (and of Eciton s. str. broadly) are
typically very large. In all probability the broods in this season
comprise more than 25,000 individuals as a rule. That a single
queen is capable of delivering an egg mass of these proportions
within a few days is an impressive fact.

Beyond the fact that in nearly twenty captures we have never
found more than one queen in a colony, and the fact that the
enormous clutch of eggs in colony 38 H may be attributed safely
to a single queen, our evidence on condition of the broods seems
to exclude any possibility that more than one queen figures in
producing a given brood. First of all, the range and central
tendency of body size in young and advanced larval broods are
rather constant for given stages (see Table I). The range of
individual sizes is at first limited ( e.g colonies 38 H, A, B, and
X) but increases with the general age of the brood (colonies 38 C,
D, E , F, G, and others). That there is a definite unimodality
in the distribution of individual sizes within a given brood was
indicated by study of the broods mentioned above, and has been
verified in all four of the broods surveyed in toto. Furthermore,
all brood distributions are skewed toward the worker-minor ex-
treme. With the population classified into five body-size cate-
gories (in terms of body length), with No. 1 the smallest and No.
5 the largest, the mode falls in category No. 2 as a rule, well below
the median value. For example, in a hamatum pupal brood (of
31,379 individuals in cocoons) which was surveyed, the following
distribution was found through the five size categories, from
smallest (ca. 5 mm. long) to largest (ca. 10.5 mm. long) : 5,209,
16,860, 8,072, 870, and 368, respectively. The prevalence of the
described distribution pattern in Eciton brood populations indi-
cates the involvement of a single reproductive source in each case,
a maternal source having definite and regular properties.

As further evidence we may repeat the fact that enclosure of
the larval brood and the opening of cocoons when a pupal brood
hatches are matters of mass change, both events accompanied
by prominent changes in colony behavior. In view of these con-
siderations together with our invariable discovery of but one
queen to a colony in many captures, the conclusion seems justified

June, 1944]

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185

that these large worker broods appearing in hamatum (and in
burchelli) colonies are attributable to single queens in the respec-
tive cases.17

Furthermore, the time relations of successive broods in given
colonies are sufficiently predictable to suggest the function of but
one queen in each case. The results from colony 36 A (E. ha-
matum), which was studied during a period of 42 days, from
August 5 to September 15, 1936, bear directly on this point
(Schneirla, 1938). During seven weeks of observation, this
colony passed from a statary period into a nomadic period which
lasted 17 days, then spent 19 days in a new statary period, after
which it entered a further nomadic period. Three successive
broods were observed, all of them relatively immense and entirely
composed of worker forms. The first of these broods terminated
its development and appeared as callow workers when the study
began. The second brood was present as developing larvae
through the complete nomadic period, was enclosed in cocoons
and entered the phase of pupation when the fully observed statary
period began, passed through its pupal development and ap-
peared as callows as a further nomadic period got under way.
The eggs of the third brood were laid at some time between Au-
gust 28 and September 6, and this brood was well started on its
larval development when the colony was captured September 15.
In the end the colony was anaesthetized and thoroughly examined.
Only one queen (in the contracted condition) was found.

This evidence is brought together in the schematic representa-
tion of the hamatum behavior cycle in Figure 2. The condition
of the brood or broods as indicated at times of major change in
colony behavior corresponds to the facts for colony 36 A and
checks with many other cases, as does the representation of a
developing larval brood through the nomadic period and a pupal
brood through the statary period. Our records indicate that in
the area of this study individual hamatum (and burchelli) col-
onies characteristically pass through the cycle of correlated be-
havior changes and worker-brood production a number of times

17 A possible alternative explanation is that the broods are the product of
two or more queens with exactly synchronized visceral rhythms. This seems,
very doubtful, especially because no polygynous colonies have been dis-
covered.

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in regular succession during the first four or five months of the
rainy season.

To describe the characteristic time relations of the principal
changes in brood development for hamatum, it is necessary to
ascertain the time at which the eggs are laid more exactly than is
possible from the record of colony 36 A. From general evidence,
and from the facts concerning queen 38 H, we have inferred that
the process of egg-laying occurs during a short period following
about one week after the beginning of the statary period. The
conclusion that normally a new brood appears as eggs about ten
days before the end of each given statary period is supported
directly and indirectly by numerous facts. In particular, al-
though eggs are never found in colonies examined during the first
few days of the statary period, large masses of eggs are found
during the last week of this period. Queen 38 H evidently had
just passed the peak of an egg-laying process when captured seven
days after the beginning of a statary period. Since in hamatum
oviposition appears to be of short duration, probably no more
than three or four days from beginning to end, if we set 18-20
days as the usual length of the statary period in this species we
may say that the new brood has roughly ten days of early develop-
ment before the statary period ends.

On this basis we may estimate the developmental period of a
given brood. To the first 10 days of embryonic and early larval
growth when the colony is statary, and 17 days to complete larval
development in the nomadic period, we may add 19 days (as a
close approximation) in the pupal stage during the following
statary period. That makes a total of 46 days for the complete
development of a hamatum brood. Other facts plainly indicate
that the interval between the appearance of successive broods in
a given colony is the number of days from the end of one statary
period (when one brood is delivered as callows) to the end of the
next statary period (when the next brood is delivered), i.e., about
35 days.18

is As may be seen in Figure 2, the difference of about 10 days in the above
figures is attributable to the fact that the oviposition and early development
of a given new brood overlaps the period in which the preceding brood is
completing its pupal maturation.

June, 1944]

Schneirla: Insect Behavior

187

On this basis we may arrive at an approximation of the interval
between successive occurrences of the short but strenuous ovi-
position act of the queen. After having deposited a given clutch
of eggs, the queen evidently lays no more until the early part of
the next statary period. If the peak of one process is roughly
placed at seven days after the beginning of a statary period
and its end at nine days, for the ensuing resting phase there are
ten days remaining in the same period, 17 days in the ensuing
nomadic period, and perhaps five days in the following statary
period, — 32 days in all for the interval between successive inter-
vals of actual egg-laying. This checks fairly well with the figure
offered above for the interval between the appearance of suc-
cessive broods of callows, as it obviously should.

To repeat, there is every reason to believe that in a given
hamatum colony the broods appearing at intervals of about 35
days are the progeny of a single queen. Not only the facts con-
cerning the capture of queens, but also the highly predictable
time relations between successive broods, support this conclusion.
The facts as sketched in Figure 2 indicate that the entire set
of events is highly synchronized; particularly in the ability of
the queen to deliver a new batch of eggs about 12 days before
the previous brood joins the worker population of the colony.
The evidence suggests that in our area of study a given hamatum
queen is capable of repeating this process a number of times
during the first months of the rainy season and perhaps even
longer.

Thus in an indirect manner but nevertheless very effectively
the queen is the pace-setter of the cyclic changes which have
been described for the type species. This relationship appears
to hold for other Eciton s. str. species as well ( e.g E. burchelli,
E. vagans), and perhaps also in other Eciton subgenera.

To summarize, in hamatum we seem to have the master pattern,
as it were : 1 ) the queen at regular intervals and in a strikingly
precise manner furnishes a huge mass of eggs which begin their
development almost simultaneously, 2) the given brood does not
materially influence events until the emergence of the previous
brood as callows dynamizes the population into nomadism, 3)
then the new brood (as larvae) takes over as principal source of

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the “social stimulation” which maintains the highly dynamic
activity pattern of the colony for some time ; 4) when the “drive”
effect of the larval brood is removed as it enters the pupal condi-
tion the colony lapses into its minimal activity pattern, from
which 5) the emergence of this brood as callow workers arouses
it into a new dynamic phase — and so on.

The queen is the key of this entire process, but only in an
indirect manner through her physiological properties as the pro-
ducer of successive broods. After a given brood has appeared as
eggs we may say hypothetically that the presence of the queen
is essential neither for the next major change in colony behavior
nor for the maintenance of this change ( i.e ., for nomadism). In
fact without the queen the next sessile phase of the colony-
behavior cycle may also occur and the colony will even begin
its next nomadic period (c/. Fig. 2). However, if no new brood
is forthcoming we should expect the colony to lapse from its new
nomadic phase into an aberrant and mainly sessile type of ex-
istence, i.e., a protracted statary condition. Some evidence
corroborating this prediction is in fact available from the study
of colonies deprived of their broods.

To state the hypothesis in a different way, it is quite likely
that in the dry season when Eciton broods are very small (either
through metabolic insufficiency in the queen or through brood
cannibalism in the workers, or both of these), the colonies lapse
almost completely into statary life. But under optimal condi-
tions in the rainy season, the adequate condition and regular
labors of the queen periodically furnish the colony, in an inci-
dental manner, with the changes in trophallaxis-based social
stimulation which condition the drive (or lack of drive) under-
lying alternate changes in the general behavior cycle.

The exquisite synchronization of the set of relationships in-
volved in the Eciton behavior pattern is emphasized in several
ways when the queen is considered in her role of key individual.
Through her capacity to deliver an entire batch of eggs within
a few days, the queen indirectly contributes the precision with
which the colony shifts from one mode of life to another. This in
turn promotes a further characteristic of great adaptive value.
Due to the spacing of her successive broods, the queen becomes

June, 1944]

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189

physogastric and lays her eggs only at times when the colony
happens to be statary (see Fig. 2), a combination of events which
is most opportune for survival of colony and species. It is ap-
parent from our discussion of the indirect relations existing
between the cycles of queen and colony functions that a true
convergence of events occurs in this case, which serves to guard
the queen from injury at the one time she is most vulnerable.
When she is gravid, having to make her way over a long bivouac-
change trek undoubtedly would offer a serious risk for the queen,
carrying the constant threat of dangers such as a tumble from
elevated sections of the path on vines or tearing the tightly
stretched intersegmental membranes on rough surfaces. In-
stead, thanks indirectly to her own visceral regularity, the queen
is safely immured through this critical period, and moreover,
there follows a further time of safety during which she may
convalesce from the organic ordeal of large-scale oviposition.
Not least among the factors fitting into the marked adaptivity of
this synchronized pattern of events, the bivouac of the colony in
its statary phase is characteristically more sequestered than at
other times. Altogether, it would be difficult to find a more
effectively adaptive mosaic of various biological events than this
one, based upon the physiological properties of the Eciton queen.

SUMMABY AND CONCLUSIONS

The functions of the Eciton hamatum queen serve indirectly
as a pace-making factor in the colony behavior pattern of the
species.

The determining processes in a given colony center around the
reproductive properties of the single functional queen. Except
when colony bivouac-change movements occur the fertile queen
is confined to the bivouac, evidently to a large extent because of
her pronounced photonegativity. At regular intervals of ap-
proximately 35 days the dichthadiigyne becomes physogastric and
within a few days releases a mass of more than 20,000 eggs. Be-
tween egg-delivery episodes she remains in the contracted or
“resting’’ condition. The ability of the hamatum queen to
mature and deliver an immense number of eggs within a very
limited time means that all members of a given brood develop

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and mature roughly in synchronization. These facts hold im-
portant consequences for the colony behavior pattern.

In the rainy season of the Caribbean lower rain-forest zone of
Panama, a colony of E. hamatum passes through statary (mini-
mal raiding; absence of migration) and nomadic (maximal daily
raiding; successive daily bivouac-changes) behavior phases at
regular intervals, each period lasting nearly 20 days. Depending
upon the properties of a fertile queen, the alternation of these
behavioral phases is highly predictable.

Significantly different intracolony circumstances parallel the
two phases of the colony behavior cycle. Colonies in the nomadic
condition invariably contain a single brood passing through its
larval stage, and the period ends precisely when this brood has
become mature and is mainly enclosed in cocoons. In statary
colonies a brood in the pupal stage is found, and the period ends
when this brood has matured and has mainly emerged as callow
workers. Approximately seven days after a given statary period
has begun, a new brood of more than 20,000 eggs appears.

The intimate correspondence which exists between Eciton
colony behavior and brood condition is explicable in terms of the
Wheeler trophallaxis concept. Evidence is cited for the involve-
ment of a special social-stimulative effect, a superadded “ drive’ ’
factor, furnished by broods capable of extensive overt activity.
Emanating from a newly emerged lot of callow workers, this effect
arouses a previously sessile colony to the threshold of maximal
raiding and daily bivouac-change, and thus is responsible for
initiating a new nomadic period. Furthermore, due to a rather
precise synchronization of successive broods, before the energiz-
ing function of a new callow brood runs its course it is overlapped
and succeded by the similar function of a sufficiently developed
larval brood. This larval effect maintains the nomadic condition
over a considerable time, but the colony lapses promptly from
nomadism when the larvae mature and spin their cocoons. Thus
the sessile and relatively inactive statary condition which ensues
is attributable to the existence of an inadequate energization of
the colony when the brood (eggs; pupae) is incapable of function-
ing as an effective source of social stimulation.

Periodic changes characterizing Eciton colony behavior thus
basically depend upon a highly regular reproductive cycle in the

June, 1944]

Schneirla: Insect Behavior

191

queen. By furnishing new broods at fixed intervals, the queen’s
function indirectly governs the presence or absence of the intra-
colony dynamizing factor critically responsible for the ebb and
flow of events in the Bciton behavior pattern.

LITERATURE CITED

Andre, E. 1885. Species des Hymenopteres. (Supplement au species des
Formicides d ’Europe et des pays limitrophes.) 8: 838-840.

Bruch, C. 1934. Las formas femeninas de Eciton . . . Descripcion y re-
descripcion de algunas espeeies de la Argentina. Ann. Soc. Cient.
Argent., 118, 2nd sem., 1934: 113-135.

Emerson, A. E. 1939. Social organization and the superorganism. Amer.
Midi. Nat., 21 : 182-209.

Gallardo, A. 1920. Las hormigas de la Republica Argentina — Subfamilia
dorilinas. Anal. Mus. Nac. Hist. Nat. Buenos Aires, 30: 281-410.
Muller, W. 1886. Beobachtungen an Wanderameisen (Eciton hamatum
Fabr.).i9 Kosmos, 18: 81-93.

Reichensperger, A. 1926. Das $ von Eciton mattogrossensis Luederw.
(Hym.) Ent. Mitteil., 15: 401-404.

. 1934. Beitrag zur Kenntnis von Eciton lucanoides Em. Zool. Anz.,

106: 240-245.

Schneirla, T. C. 1933. Studies on army ants in Panama. Jour. Comp.
Psychol., 15 : 267-299.

. 1934. Raiding and other outstanding phenomena in the behavior of

army ants. Proc. Nat. Acad. Sci., 20: 316-321.

. 1938. A theory of army-ant behavior based upon the analysis of

activities in a representative species. Jour. Comp. Psychol., 25:
51-90.

. 1940. Further studies on the army-ant behavior pattern. Mass

organization in the swarm-raiders. Ibid., 29: 401-460.

. 1944. Studies on the army-ant behavior pattern. Nomadism in the

swarm-raider Eciton burchelli. Proc. Amer. Phil. Soc., Phila., 89
(1) (in press).

Weber, N. 1941. The rediscovery of the queen of Eciton ( Labidus ) coecum
Latr. (Hym.: Formicidce) . Amer. Midi. Nat., 26: 325-329.
Werringloer, Anneliese. 1932. Die Sehorgane und Sehzentren der
Dorylinen nebst Untersuchungen iiber die Facettenaugen der For-
miciden. Zeitschr. wiss. Zool., 141 : 432-524.

Wheeler, G. C. 1943. The larvae of the army ants. Ann. Entom. Soc.
Amer., 36: 319-332.

Wheeler, W. M. 1900. The female of Eciton sumachrasti Norton, with
some notes on the habits of Texas Ecitons.20 Amer. Nat., 34 : 563-
574.

19 Redetermined as E. burchelli.

20 Redetermined as E. schmitti.

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Journal New York Entomological Society

[Vol. LII

. 1910. Ants, their Structure, Development and Behavior. N. Y. :

Columbia University Press, pp. 663.

— -. 1921. Observations on army ants in British Guiana. Proc. Amer.

Acad. Arts Sci., 56 : 291-328.

. 1925. The finding of the queen of the army ant Eciton hamatum

Fabricius. Biol. Bull., 49: 139-149.

. 1928. The social insects, their origin and evolution. N. Y. : Har-

court, Brace.

and I. W. Bailey. 1920. The feeding habits of Pseudomyrminse

and other ants. Trans. Amer. Phil. Soc., 22, N.S., Art. 4: 235-279.

Zahl, P. A. 1939. To the Lost World. N. Y.: Knopf.

June, 1944]

Bird: Papaipema

193

A RE-SURVEY OF PAPAIPEMA SM. (LEPIDOPTERA)

By Henry Bird
Rye, N. Y.

The intricacies of insect life are abysmal and any individual
studies are bound to be incomplete and fragmentary. This is so
axiomatic as to need no argument and may excuse the limited
viewpoint of any single observer. However, as time goes on data
and observable facts accumulate, given problems here and there
gain enlightenment through various channels.

Retrospective deductions on the part of the writer based on
“the sum of evidence” as this slowly evolves apparently offers
some ground work for the serial arrangement of Papaipema
species.

Conceived as an ontogenetic tree with its phylogenetic roots
outcropping from supposedly more ancient genera, it is interest-
ing to invade this vale of surmise.

To what extent these relationships can be shown in a list
arrangement is unsatisfactory but should be undertaken. As
building bricks there are the characters of the adults including
of course the genitalia of both sexes, the gleaning from larval
evidence backed by attending factors. Thus it becomes requisite
to discourse somewhat at length taxonomically.

Also the final disposition of holotypes should be chronicled.

First, as to the generic basis whereupon Prof. J. B. Smith
established Papaipema in 1899. 1

He named no genotype and the gist of his characterizations
featured moths with primaries rather broad and outwardly acute
at apex; the thoracic tuftings decidedly upright and anteriorly
usually broadened, in form like an “adze” behind the collar; the
antennae are simple ; the f rons smooth ; the male genitalia mainly
show a unique pattern “having the harpes more or less forked
with triangular patch of spinulated surface at the tip. The
clasper in almost all cases a long, stout, curved hook, but is unique
in having the outer curve strongly toothed. ’ ’

1 Revision of Hydroecia, Trans. Am. Ent. Soc., Yol. XXVI.

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Journal New York Entomological Society

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As now considered there are nearly fifty species in the genus
with thirty-seven at least following this genitalic pattern closely.
Smith’s simple drawings of a portion of the male genitalia are
misleading however since it is necessary to chronicle the following
specific errors.

Harrisii and pterisii were considered by him as one species;
duovata, arctivorens and merriccata at least were confused under
“rutila”; circumlucens, ochroptena and form humuli were
treated as circumlucens ; he considered cerussata and frigida form
thalictri under cerussata label; treated purpurifascia and lysi-
machice as one; misidentified sciata for “limpida” ; nepheleptena
for appassionata, while his reference to necopina had largely to
do with maritima.

Criticism should not be levied unduly at these presumed mis-
takes since some of Guenee’s types, notably rutila and limpida,
British Museum uniques, have not been satisfactorily associated,
and two of our names as used hereinafter will probably fall
through this lack of perception.

In 1910 Sir George Hampson,2 following his custom of citing
genotypes, selected cerina as genotype of Papaipema, on First
Species Rule because it headed Smith’s enumeration of the genus.
That was an unfortunate usage since the species does not measure
fully to Smith’s definition. Recognizing this discrepancy from
a mere autopic glance, and though bound by the Rules he never-
theless uses the very proper species harrisii in illustrating vena-
tion and the bodily detail, quite representative of the genus.

Begging Sir George ’s pardon, this writer votes that cerina Grt.
be superseded as genotype of Papaipema by harrisii Grt., if a
more elastic rule be forthcoming some day.

The limitations of generic boundaries are subject to varying
personal ideas but ideally their demarcation should suggest evo-
lutionary trends in so far as that might be surmised.

Avoiding theory as much as possible but judging facts as they
appear today, we can find much aid in turning to the larvae in
their earlier stages.

It is generally conceded that early stage larvae reflect the primi-
tive ancestral line at least phylogenetically. Leaning on this

2 Cat. Lep. Phal. Brit. Mus., Vol. IX, p. 80.

June, 1944] Bird: Papaipema 195

deduction, a personal familiarity with forty Papaipema species
seems to help in some measure while details of color pattern aid
specifically. As larvae Papaipema species are unique.

Their early larval pellicle is distinctly colored whereas most
mining larvae are at all stages more or less translucent.

The great majority show a contrasting middle ring of dark
purplish or pinkish brown, in livid hue, at the first four abdomi-
nal segments while elsewhere longitudinal white or yellowish lines
drawn on the darker body color produce striking individuals.

This intensity continues through the instars up to the penulti-
mate, while maturity usually exhibits a faded translucence.

Three pattern types follow; the dark middle girdle may show
an abrupt termination of all lines ; or the dorsal line may cross it
in unbroken continuity ; or both the dorsal and subdorsal may be
entirely unbroken. These features of pattern aid much specifi-
cally.

They are constant with two exceptions — the Pacific Coast spe-
cies angelica and insulidens , where one, or rarely both lines may
be continuous. One is tempted to assume that a progenitor was
not wholly an internal feeder but subsisted within some en-
circling tissues with both extremities exposed and maintaining
there the linear markings. The above category applies to thirty-
eight known individuals. Two other known species are decidedly
different and, again assuming, feature as admirable connecting
links with their Apamea-Gortyna-Hyroecia relatives possessing
a world-wide, north temperate zone dispersal. Because of this
dispersal as against restricted North American Papaipema it
seems rational to consider the latter as a subsequent offshoot.
Their very close relationship bespeaks a comparatively recent
evolution wherein possibly marked choices of food plants, some
of the latter also restricted to America, may have played a part
in influencing specific origin.

The larvae of the two alleged connecting species, frigida and
beeriana have their markings as transverse segmental rings,
features prevailing with Apamea erepta ryensis, Hydrcecia im-
manis, H. micacea and PL. stramentosa , which is as far as famili-
arity goes.

The exotic Xanthoecia flavago larva is cross banded, while
Parapamea buffalcensis and Emboloecia sauzalitee have larvae

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which are longitudinally lined. This latter trio has the frons
distinctly armed but these various genera may all figure as part
of the ‘ ‘ Gortynid series. ’ ’ This term has been used by the writer
as a convenience, and in view of Grote’s arguments,3 Gortyna,
genotype micacea Och., might properly find generic application
somewhere therein.

As structural larval features, the heavy setigerous plates are
noteworthy, with a peculiar development in many species that
have an additional plate known as IVa, on joint ten. Since this
plate bears no setal hair its transitory character may be adduced.

However it is one of the evidences aiding specific distinction.

The rugged genitalia become a prime structural feature with
the adults. In the males such closeness to the conventional pat-
tern prevails as to indicate the very near relationship of the
species. Greatest modification exists with furcata and eryngii,
while frigida, unimoda and appassionato, make a decided break
with the harpes greatly reduced. One might suggest a generic
break here but full evidence points to connecting species.

With the female genitalia the genital plate at the ostium is
of help specifically. It is a heavy, more or less shield-shaped
process, differing in outline and scobinated characteristically.
Dr. F. Heydemann in a praiseworthy treatment of the nictitans
group of Apamea 4 notes the value of the genital plate and figures
the character without other detail upon a single plate for specific
comparison.

The close proximity of Papaipema species blend them together
into a satisfactory whole. Indeed, in a number of instances they
are so close that if no further evidence was at hand than a few
flown specimens much doubt would arise as to their distinction.

Furthermore, variation is rife particularly in the feature
wherein the ordinarily prominent white marked stigmata may
be obsolescent or vice versa. This is productive of such an autopic
difference that erroneously, two distinct species seem to be in-
volved— vide Guenee’s two alleged species, nebris and nitela.

That a varietal name be given to the lesser of such forms seems
practical. This has been done in the more striking instances and

3 Historical Sketch of Gortyna, Proc. Am. Phil. Soc., Vol. XXXIX, No. 162.

4 Die Arten der Eydroecia nictitans .« Sond. Ento. Zeit., XXXXIX u.

xxxxv.

June, 1944]

Bird: Papaipema

197

the future holds possibilities with slighter stigmatal variance
likely to be grasped by some. Some dimorphism is observable,
the ubiquitous cataphracta with its extensive food habits shows
this in its more northern range, while imperspicua represented by
a unique type may well be in that category. Until rearing proves
the fact its specific standing may remain.

Two formerly considered species must merge as one, verona
and astuta with the latter name preserved as varietal. Smith’s
unique type of verona, from Winnipeg, Manitoba, is a dwarfed
pale form of the species, while astuta was applied to a larger more
colorful variant with the terminal space solidly purplish, easily
suggestive of distinctness. Both can occur in the same locality
and though verona is less numerous in the writer’s experience;
it has priority.

That astuta be retained as a distinct color, or dimorphic form
points to future expediency.

As to the placement of Papaipema holotypes, the writer has
prepared a detailed treatment of the genus under title “The
Epic of Papaipema,” a unique copy which is willed to the
American Museum of Natural History, New York, N. Y.

With it will go his collection of the group. Hence the holotypes
and paratypes, with the relevant literature will be at one place
for future students. The collection embraces something over
fifteen hundred specimens, mainly reared. The various types
number ninety-eight examples. The aggregation brings together
not only the adults, but larval and pupal stages, to some extent
the parasites; the foodplant habitations are also shown. The
genitalic slides are not considered in this summary.

The “Epic” consists of three volumes. It assembles the
principal published literature thus dealing with the historic rise
and subsequent departures in the genus, treats monographically,
matters of taxonomic import, features of parasitism and the
gleanings of several decades of field study. Interlarded between
the published papers, the author’s notes and criticisms bring
such up to current review. Particularly some of his earlier
papers were rather puerile and in need of revision. Volumes I
and II total 1127 pages, inclusive of the relevant articles. Vol-
ume III is a portmanteu affair.

198

Journal New York Entomological Society

[Vol. LII

Seventy plates occupying seven containers, are unbound for
easy comparison; four containers hold autographed letters from
important workers in connection with the subject, many of whom
have now passed on. The plates illustrate by line drawings the
infested food plants in some instances, larval features and the
genitalia of both sexes in so far as possible, while the adults are
shown in color. There is also a booklet of remarks and index of
plates.

By what manner a commingling of the species of Papaipema
can be best portrayed with their ontogenetic proximity appar-
ently shown, it is convenient to resort to a popular vegetative
process and erect a fanciful “tree.”5 Then, as nonconventional
genes seem to have effected the protoplasmic stream, branches or
shoots may materialize, either ascending or paralleling the main
trunk according to the line of thought. Admirable as a basis of
ideas, but to transplant this fruitage to the linear order of list
column spoils the conception entirely. The writer can only sug-
gest the following summary.

List order for the species of the genus

PAPAIPEMA Smith.

(Asterisk denotes larva unknown, synonyms in italics)

frigida Smith.

form thalictri Lyman.
terminalis Strand,
unimoda Smith.*
beeriana Bird.

form lacinarise Bird,
appassionata Harvey.
horni Strand.

purpurifascia Grote & Robin-
son.

luteipicta Strand,
lysimachiae Bird.

nec purpurifascia Auct.
stenoscelis Dyar.
speciosissima G. & R.

form regalis Wyatt & Beer,
s EPIC OF PAPAIPEMA, 1940, pp.

inquaesita G. & R.
form wyatti Barnes & Ben-
jamin,
pterisii Bird.

triorthia Dyar.
anargyria Dyar.*
ochroptena Dyar.

form humuli Bird,
arctivorens Hampson.
merriccata Bird,
araliae Bird & Jones,
harrisii Grote.

form mulieris Strand,
sub. sp. rubiginosa Bird,
verona Smith.

form astuta Bird.

553-554, Yol. II.

June, 1944]

Bird: Papaipema

199

rutila Guenee.*
depictata Benjamin.*
nepheleptena Dyar.

moeseri Bird,
impecuniosa Grote.
circumlucens Smith.
baptism Bird,
form ochroptenoides
Benj.*

sub. sp. vaha Benj.*
marginidens Guenee.
birdi Dyar.

nephrasyntheta Dyar.*
furcata Smith,
rigida Grote.
pertincta Dyar.
limata Bird*
insulidens Bird,
angelica Smith,
cataphracta Grote.
form sulphurata Bird,
race fluxa Bird,
imperspicua Bird.*

duovata Bird,
aerata Lyman,
placida Bird.*
cerina Grote.
dribi Benjamin.*
polymniae Bird,
nebris Guenee.

form nitela Guenee.
duplicata Bird.

obsolescens Strand,
silphii Bird,
necopina Grote.
nelita Strecker.
form linda Bird,
form obicularis Strand,
errans Barnes & McDunnough.
engelhardti Bird,
sciata Bird,
limpida Guenee.*
cerussata Grote.
eryngii Bird,
maritima Bird,
eupatorii Lyman.

200

Journal New York Entomological Society

[Vol. LII

INSECT INTRODUCTIONS AND WAR

The recent press releases on the introduction of potential insect
pests in packages sent home by members of the armed forces over-
seas has evidently stirred some interest. The Bureau of Plant
Industry, New Jersey Department of Agriculture, was called to
inspect and fumigate a reed stool received by a woman in Trenton,
New Jersey from her son in India. This was found to be infested
with numerous small bostrichid beetles which were identified by
W. S. Fisher, Bureau of Entomology and Plant Quarantine at
Washington, D. C., as Dinoderus brevis Horn. D. brevis was
originally described from a single specimen taken in Louisiana,
and was believed to be a native species. This species was later
found to be Oriental, especially common in India, and has been
carried, through commerce, to many parts of the world. — William
M. Boyd.

June, 1944]

Rapp: Psychodid^e

201

CATALOGUE OF NORTH AMERICAN PSYCHODID^

By William F. Rapp, Jr.

In 1905 Aldrich1 published the last catalogue of North Amer-
ican Diptera. In the family Psychodidee he listed 34 species,
which were distributed in four genera. Table I is a comparison
of Aldrich’s 1905 catalogue and this catalogue.

TABLE I

Genus Aldrich 1905 Rapp 1943

Flebotomus 3

Pericoma .. .... 11 13

Psychoda 21 41

Sycorax 1

Trichomyia 1 2

No. of species 34 59

Since 1905 several workers have been attracted to this family
with the result that many changes have arisen in nomenclature,
plus the addition of new species. This catalogue has been com-
piled after a careful survey of the entomological literature.
Synonymy is listed wherever it has appeared in the literature.
It is interesting to note that of the 34 species listed by Aldrich
only 6 have been reduced to synonymy. Of the 54 species de-
scribed since 1905, 19 have proven to be synonyms.

The North American Psychodidae, as a whole, are of little eco-
nomic importance. Certain species of Flebotomus are known
vectors of tropical diseases, but none of these occur in the North
American region. Psychoda alternata Say is at times a pest
around sewage filter plants.2

The area covered in this paper includes all of America north
of Mexico, or the area considered as the nearctic region based
upon zoogeographical division.

1 Aldrich, John M. “A Catalogue of North American Diptera,” Smith-
sonian Miscellaneous Collections , Yol. 46 (1905), p. 1-680.

2 Headlee, T. J., and Beckwith, C. S., ‘ ‘ Sprinkling Sewage Ply, Psychoda
alternata Jour. Econ. Ent., Yol. 11 (1918), p. 395-401.

202

Journal New York Entomological Society

[Vol. LII

FLEBOTOMUS* Rondani
Flebotomus diabolicus Hall.

Phlebotomies diabolicus Hall, Proc. Ent. Soc. Wash., Vol. 38
(1936), p. 28.

Texas.

Flebotomus texanus Dampf.

Phlebotomies texanus Dampf, Anales de la Escuela Nacional
de Ciencias Biologicas, Vol. 1 (1938), pp. 119-122.
Texas.

Flebotomus vexator Coquillett.

F. vexator Coquillett, Ent. News, Vol. 18 (1907), p. 102.
Louisiana, Maryland.

PERICOMA Walker
Pericoma bipunctata Kincaid.

P. bipunctata Kincaid, Ent. News, Vol. 10 (1899), p. 34.
California, Washington.

Pericoma californica Kincaid.

P. californica Kincaid, Ent. News, Vol. 12 (1901), p. 195.
California.

Pericoma Carolina Banks.

P. Carolina Banks, Bui. Brooklyn Ent. Soc., Vol. 26 (1931),

p. 228.

North Carolina.

Pericoma furcata Kincaid.

P. furcata Kincaid, Ent. News, Vol. 10 (1899), p. 34.
Washington.

Pericoma longiplata Haseman.

P. longiplata Haseman, Trans. Am. Ent. Soc., Vol. 33 (1907),
p. 308.

Arizona.

Pericoma ocellaria var. americana Kincaid.

P. ocellaria var. americana Kincaid, Ent. News, Vol. 12
(1901), p. 194.

Maine.

Pericoma satellitia Dyar.

P. satellitia Dyar, Proc. Ent. Soc. Wash., Vol. 29 (1927),
p. 163.

Maryland.

* Phlebotmus of authors.

June, 1944]

Rapp: Psychodidjs

203

Pericoma scala Haseman.

P. scala Haseman, Trans. Am. Ent. Soc., Vol. 33 (1907),
p. 307.

Arizona.

Pericoma sitchana Kincaid.

P. sitchana Kincaid, Ent. News, Vol. 10 (1899), p. 33.
Alaska, Oregon.

Pericoma trialbawhorla Haseman.

P. trialbawhorla Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 306.

Missouri.

Pericoma triloba Kincaid.

P . triloba Kincaid, Ent. News, Vol. 10 (1899), p. 33.
Washington.

Pericoma truncata Kincaid.

P. truncata Kincaid, Ent. News, Vol. 10 (1899), p. 35.
California.

Pericoma variegata Kincaid.

P. variegata Kincaid, Ent. News, Vol. 10 (1899), p. 33.
Washington.

PSYCHODA Latreille
Psychoda alberta Curran.

P. alberta Curran, Can. Ent., Vol. 56 (1924), p. 219.

Alberta.

Psychoda albipunctata Williston.

P. albipunctata Williston, Ent. News, Vol. 5 (1893), p. 113.
Telmatoscopus meridionalis Eaton, Ent. Mo. Mag. (1894), p.
195.

P. snowii Haseman, Trans. Am. Ent. Soc., Vol. 33 (1907), p.
311-312.

P. erect a Curran, Cat. Ins. Jam. Dept. Agr., Jamaica Ent.

Bui. No. 4, pt. 1 & 2 (1926), p. 102.

Florida, Louisiana, South Carolina, Texas.

Psychoda albitarsis Banks.

P. albitarsis Banks, Can. Ent., Vol. 27 (1895), p. 324.
Maryland, New Jersey, New York, North Carolina, Virginia.
Quebec.

204

Journal New York Entomological Society

[Vol. LII

Psychoda alternata Say.

P. alternata Say, Long’s Exp. St. Peter’s River, App. (1824),
p. 358.

Tipula phalcemoides Scop., Ent. Carn., No. 864 (1763), p.
324.

Psychoda sexpunctata Cnrtis, Brit. Ent., Vol. 16 (1839), p.
745.

P. marginepunctata Roser., Corr. Wurt. landro, Ver. 1
(1840), p. 50.

P. schizura Kincaid, Ent. News, 10 (1899), p. 32.

P. floridica Haseman, Trans. Am. Ent. Soc., 33 (1907), p. 316.
P. nocturnala Haseman, Trans. Am. Ent. Soc., 33 (1907), p.
319.

P. bengalensis Brunetti, Rec. Ind. Mus., Vol. 11 (1908), p.
370.

P. albimaculata Welch, Ann. Ent. Soc. Amer., Vol. 5 (1912),
p. 411.

P. dakotensis Dyar, Insec. Inscit. Menst., Vol. 14 (1926), pp.
107-110.

California, Connecticut, District of Columbia, Florida, Illi-
nois, Kansas, Maryland, Missouri, New Hampshire, New
Jersey, New Mexico, New York, Ohio, Oregon, Pennsylvania,
South Dakota, Texas, Virginia, Washington.

Psychoda annulipes Johnson.

P. annulipes Johnson, Bui. Amer. Mus. Nat. History, Vol. 32
(1913), p. 43.

Florida.

Psychoda aterrima Banks.

P. aterrima Banks, Ent. News, Vol. 25 (1914), p. 128.

New York.

Psychoda augusta Curran.

P. augusta Curran, Can. Ent., Vol. 58 (1926), p. 228.

Quebec.

Psychoda autumnalis Banks.

P. autumnalis Banks, Ent. News, Vol. 25 (1914), p. 127.
Pericoma litt oralis Dyar, Insec. Inscit. Menst., Vol. 14
(1926), pp. 107-110.

Pericoma aldrichana Dyar, Insec. Inscit. Menst., Vol. 14
(1926), pp. 107-110.

June, 1944]

Kapp: Psychodid^e

205

California, District of Columbia, Maryland, Virginia.
Alaska.

Psychoda bicolor Banks.

P. bicolor Banks, Can. Ent., Vol. 26 (1894), p. 33.

P. nigra (Banks) Dyar, Proc. Ent. Soc. Wash., Vol. 30
(1928), p. 87.

District of Columbia, Indiana, Maryland, New York.
Psychoda bishoppi Del Rosario.

P. bishoppi Del Rosario, Philippine Jour. Sci., Vol. 59 (1936),
p. 141.

Maryland.

Psychoda cinerea Banks.

P. cinerea Banks, Can. Ent., Vol. 26 (1894), p. 331.

P. elegans Kincaid, Ent. News, Vol. 8 (1897), p. 144.
Threticus compar Eaton, Ent. Mo. Mag. II, Vol. 15 (1904),
p. 57.

P. domestica Haseman, Ent. News, Vol. 19 (1908), p. 285.

P. compar Tonnoir, Ann. Soc. Ent. Belg., Vol. 62 (1922), p.

67.

P. prudens Curran, Can. Ent., Vol. 56 (1924), p. 219.
Connecticut, District of Columbia, Indiana, Maine, Mary-
land, Massachusetts, Missouri, New Jersey, New York, Ore-
gon, Virginia, Washington.

Alberta.

Psychoda criddlei Curran.

P. criddle Curran, Can. Ent., Vol. 56 (1924), p. 218.

Ontario.

Psychoda degenera Walker.

P. degenera Walker, List of the Specimens of Dipterous In-
sects in the Collection of the British Museum, List I
(1848), p. 33.

Ontario.

Psychoda helicis Dyar.

P. helicis Dyar, Proc. Ent. Soc. Wash., Vol. 31 (1929), p. 63.
Maryland.

Psychoda horizontala Haseman.

P. horizontala Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 313.

Missouri.

206

Journal New York Entomological Society

[Vol. LII

Psychoda interdicta Dyar.

P. interdicta Dyar, Proc. Ent. Soc. Wash., Vol. 30 (1928),

p. 88.

Maryland, New York.

Psychoda interrupta Banks.

P. interrupta Banks, Proc. Ent. Soc. Wash., Vol. 8 (1907),
p. 150.

Maryland.

Psychoda juno Curran.

P. juno Curran, Can. Ent., Vol. 58 (1926), p. 228.

Ontario.

Psychoda longifringa Haseman.

P. longifringa Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 318.

Florida.

Psychoda marylandana Del Rosario.

P. marylandana , Del Rosario, Philippine Jour. Sci., Vol. 59
(1936), p. 111.

Maryland.

Psychoda megantica Curran.

P. megantica Curran, Can. Ent., Vol. 56 (1924), p. 217.
Quebec.

Psychoda minuta Banks.

P. minuta Banks, Can. Ent., Vol. 26 (1894), p. 331.
Connecticut, New Jersey, New Mexico, New York, Virginia.

Psychoda nigra Banks.

P. nigra Banks, Can. Ent., Vol. 26 (1894), p. 331.

P. marginalis Banks, Can. Ent., Vol. 26 (1894), p. 333.
P.apicalis Banks, Proc. Ent. Soc. Wash., Vol. 8 (1906), p.
148.

P. basalis Banks, Proc. Ent. Soc. Wash., Vol. 8 (1906), p. 149.
Pericoma orillia Curran, Can. Ent., Vol. 56 (1924), p. 218.

P. varitarsis Curran, Can. Ent., Vol. 56 (1924), p. 220.
Maruina nigra (Banks) Dyar, Insec. Inscit. Menst., Vol. 14
(1926), p. 111.

Pericoma apicalis (Banks) Dyar, Insec. Inscit. Menst., Vol.
14 (1926), p. 149.

Connecticut, District of Columbia, Indiana, Maine, Mary-

June, 1944]

Rapp: Psychodid^:

207

land, New Hampshire, New Jersey, New York, Ohio, Vir-
ginia.

Ontario, Quebec.

Psychoda nitida Banks.

P. nitida Banks, Can. Ent., Vol. 33 (1901), p. 275.

District of Columbia, New York.

Psychoda olympia Kincaid.

P. olympia Kincaid, Ent. News, Vol. 8 (1899), p. 144.
Pericoma olympia (Kincaid) Haseman, Trans. Am. Ent.

Soc., Vol. 33 (1907), p. 305.

Washington.

Psychoda opposata Banks.

P. opposata Banks, Can. Ent., Vol. 33 (1901), p. 274.
District of Columbia, Maryland, New York.

Psychoda phalaemoides (Linn.) Tonnoir.

Tipula phalcenoides Linnaeus, Syst. Nat. ed. 10, No. 32
(1758), p. 588.

P . phalcenoides (Linn.) Tonnoir, Ann. Soc. Ent. Belg., Vol.
62 (1922), p. 67.

P. pacifica Kincaid, Ent. News, Vol. 8 (1897), p. 143.

P. tonnoiri Dyar, Insec. Inscit. Menst., Vol. 14 (1926), p. 105.
California, Colorado, District of Columbia, Idaho, Maryland,
New Mexico, Oregon, Washington, Wisconsin.

Alberta, British Columbia.

Alaska.

Psychoda pusilla Tonnoir.

P. pusilla Tonnoir, Ann. Soc. Ent. Belg., Vol. 62 (1922),
p. 83.

Kansas, Maryland.

Psychoda quadripunctata Banks.

P. quadripunctata Banks, Ent. Soc. of Wash., Vol. 8 (1907),
p. 149.

Virginia.

Psychoda scotiae Curran.

P. scoticB Curran, Can. Ent., Vol. 56 (1924), p. 216.

Nova Scotia, Quebec.

Psychoda severini Tonnoir.

P. severini Tonnoir, Ann. Soc. Ent. Belg., Vol. 62 (1922),
p. 78.

208

Journal New York Entomological Society

[Vol. Lll

California, District of Columbia, Maryland, Montana, New
Mexico.

British Columbia.

Psychoda sigma Kincaid.

P. sigma Kincaid, Ent. News, Vol. 10 (1901), p. 31.
P.surcoufi Tonnoir, Ann. Soc. Ent. Belg., Vol. 62 (1922),
p. 74.

Washington.

Psychoda signata Banks.

P. signata Banks, Can. Ent., Vol. 33 (1901), p. 274.

District of Columbia, Maine.

Psychoda slossoni Williston.

P. slossoni Williston, Ent. News, Vol. 4 (1893), p. 114.

Maine, New York.

Psychoda snowhilli Del Rosario.

P. snowhilli Del Rosario, Philippine Jour. Sci., Vol. 59
(1936), p. 140.

Maryland.

Psychoda squamosa Johnson.

P. squamosa Johnson, Bui. Amer. Mus. Nat. Hist., Vol. 32
(1913), p. 43.

Psychoda superba Banks.

P. superba Banks, Can. Ent., Vol. 26 (1894), p. 332.

District of Columbia, Maryland, Michigan, New Jersey, New
York, Virginia.

Psychoda superba var. conspicua Del Rosario.

P. superba var. conspicua Del Rosario, Philippine Jour. Sci.,
Vol. 59 (1936), p. 125.

Maryland, Virginia.

Psychoda tridactila Kincaid.

tridactila Kincaid, Ent. News, Vol. 19 (1899), p. 32.
Washington.

Psychoda uniformata Haseman.

P. uniformata Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 319.

Missouri.

June, 1944]

Rapp: Psychodid^e

209

Psychoda uniformis Del Rosario.

P. uniformis Del Rosario, Philippine Jonr. Sci., 59 (1936),
p. 113.

Maryland.

TRICHOMYIA Haliday

Trichomyia lanceolata Kincaid.

Sycorax lanceolata Kincaid, Ent. News, Vol. 10 (1899), p. 35.
California, Washington.

Trichomyia unipunctata Haseman.

T. unipunctata Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 323.

Arizona.

June, 1944]

Heming: International Commission

211

RECENT WORK BY THE INTERNATIONAL COMMIS-
SION ON ZOOLOGICAL NOMENCLATURE

By Francis Hemming

Secretary to the International Commission on Zoological
Nomenclature

The International Commission on Zoological Nomenclature are
now engaged in the publication of decisions taken before the out-
break of war in 1939. These decisions have been embodied in
Opinions 134^183 and Declarations 10-12. Of these, Opinions
134-155 and the 3 Declarations have already been published or
are in the press. The remainder are ready for printing and will
be published as soon as funds are available.

These Opinions are of particular interest to entomologists, since
in addition to 9 Opinions relating to the interpretation of various
aspects of the International Code, no less than 38 of these Opinions
are directly concerned with entomological subjects.

The Opinions relating to the interpretation of the Code are :
Opinion 138 (meaning of phrase “definite bibliographic refer-
ence” in Article 25) ; Opinion 141 (naming of families) ; Opinion
145 (status of names first published in invalid works) ; Opinion
147 (generic names of same origin and meaning as older generic
names) ; Opinion 148 (status relating to names published as
emendations of, or substitutes for, older names) ; Opinion 164
(position as regards types when two or more genera are united) ;
Opinion 168 (supplementing Opinion 65 regarding genera based
upon erroneously determined species) ; Opinion 172 (status of
type — designations of genera in abstracts, etc.) ; and Opinion 183
(interpretation of Article 8 relating to form in which generic
names should be published).

Of the Opinions specially concerned with entomological sub-
jects, 6 deal with the status of particular works or with the dates
of such works : Opinion 136 (Latreille, 1810) affects all Orders
of insects; Opinion 135 (“Erlangen List,” 1801) is of special in-
terest to hymenopterists ; Opinions 134 (Freyer, Neue Beitrage),
138 (Hiibner, Samml. exet. Sckmett 1807 and Fabricius Mag.

212

Journal New York Entomological Society

[You Lll

Insektenk. (Illiger) 1807) and 150 (Hiibner, Verz. bek. Schmett)
are concerned with Lepidoptera and Opinion 152 (Meigen, 1800)
with Diptera. Opinions 140 and 143 deal with certain family
names in insects.

The remaining 30 Opinions deal with particular generic names
in various Orders of insects. These Opinions either fix the types
of these genera or add the names to the Official List of Generic
Names in Zoology, or do both. Three of these Opinions are con-
cerned with Orthoptera; 13 with Hymenoptera and 14 with
Lepidoptera.

Bach Opinion is published separately but Opinions are con-
secutively paged to facilitate the publication of an index on the
completion of the volume concerned.

The International Commission are most anxious to secure that
Opinions are published as rapidly as possible but they are greatly
hampered by lack of funds. The Commission therefore appeal to
scientific institutions and individual scientific workers for dona-
tions to a special fund to be used for the issue of publications.
Full particulars of this Appeal are given in Part 2 of the Com-
mission’s Official Organ, the Bulletin of Zoological Nomenclature
published in 1943.

Contributions, however small, will be warmly welcomed and
will be acknowledged in the Bulletin. Bankers’ drafts, cheques,
and money orders should be made payable to the ‘ ‘ International
Commission on Zoological Nomenclature” and sent to the Com-
mission at their Publications Office, 41 Queen’s Gate, London,
S.W.7. All orders for the Commission’s publications should be
sent to the same address. Inquiries relating to the work of the
Commission should be addressed to me at 83 Fellows Road (Gar-
den Flat), London, N.W.3.

International Commission on

Zoological Nomenclature,

Publications Office,

41 Queen ’s Gate,

London, S.W.7.

24th April, 1944.

VoL LII

No. 3

Edited by HARRY B. WEISS

1AL

Publication Committee

HARRY B,

SEPTEMBER, 1944

Journal

of the

New York Entomological Society

Devoted to Entomology in General

WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.
LANCASTER, PA.

NEW YORK, N. Y.

1944

CONTENTS

The Remarkable Distribution of an American Cicada:

A New Genus and Other Cicada Notes

By William T. Davis 213

Ephraim Porter Felt

By Stanley W. Bromley 223

Notes on Mexican Butterflies, III

By F. Martin Brown 237

Outdoor Protection from Mosquitoes

By Joseph M. Ginsburg 247

New Neotropical Theclinae (Lepidoptera, Lycaenidae)

By Harry K. Clench 255

Note on the Death-Feint of Bruchus obtectus Say

By Harry B. Weiss 262

Differentiation of Females of Certain Species of Culex
by the Cibarial Armature

By Charles D. Michener 263

Insect Responses to Colors

By Harry B. Weiss 267

Hibernation of the Syrphid Fly, Lathyrophthalmus seneus
Scop.

By E. Gorton Linsley 272

Two New Subspecies of Lycaenopsis pseudargiolus Bdv.

& Lee. (Lepidoptera, Lycaenidae)

By Harry K. Clench 273

New Species of Neopasites with Notes Concerning
Others (Hymenoptera, Nomadidae)

By E. Gorton Linsley 277

The Death-Feints of Alobates pennsylvanica DeG., and
Alobates barbata Knoch. (Coleoptera)

By Harry B. Weiss 281

Insect Food Habit Ratios of New York State

By Harry B. Weiss 283

NOTICE : Volume LII, Number 2, of the Journal of The
New York Entomological Society was published on
July 5, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.
under the Act of August 24, 1912

Acceptance for mailing at special fate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LII September, 1944 No. 3

THE REMARKABLE DISTRIBUTION OF AN
AMERICAN CICADA; A NEW GENUS,

AND OTHER CICADA NOTES1

By William T. Davis
Staten Island, N. Y.

QUESADA GIGAS, A REMARKABLE CICADA
(Plate VIII, Figs. 1, 2)

A number of species of North American cicadas occur from
the Atlantic seaboard to about western Kansas, or to the one
hundredth meridian, and northward from Florida and Southern
Texas to Nova Scotia and the region of the Great Lakes. In
Western North America, where there are a greater number of
' species than in the Eastern States and Canada, the species extend
eastward to about the 100th meridian. Two species of Okana-
gana, namely rimosa and canadensis , in the northern part of their
range, where they follow the belt of evergreen forest extending
from Nova Scotia to the far west, have a greater east and west
distribution than the cicadas occurring more to the south. In:
“A Preliminary Review of the West Coast Cicadidse,” 1915, Mr.
E. P. Van Duzee states regarding rimosa: “This species occurs
across the whole continent from Vancouver Island to Quebec
and as far south on the west coast as Fresno Co., California.”

In Texas there are at least 40 species of cicadas and several
additional named forms or varieties. The most famous species
occurring in Texas, is Quesada gigas Olivier, remarkable on
account of its distribution as well as its loud song, which has been
1 1 am indebted to Howard Cleaves for photographing the specimens.

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likened by many observers to the shrill whistle of a first-rate
locomotive. In: “The Naturalist on the Eiver Amazons,” Bates
states: “Added to these noises were the songs of strange cicadas,
one large kind perched high on the trees around our little haven
setting up a most piercing chirrup ; it began with the usual harsh
jarring tone of its tribe, but this gradually and rapidly became
shriller, until it ended in a long and loud note resembling the
steam-whistle of a locomotive engine. Half-a-dozen of these
wonderful performers made a considerable item in the evening
concert. I had heard the same species before at Para, but it
was there very uncommon. ’ ’

The common name for gigas in some localities is the: “Loco-
motive Cicada,” and it is also called the: “Soupbug,” because
it is at times attracted to light in its evening flights and lands on
the supper table.

It is the only known American cicada that may be found in
the mature or winged form every month in the year in some part
of its extensive north and south distribution of about 4,000 miles,
from southern Texas through Mexico, Central America, South
America into Argentina.

In the writer’s collection there are many specimens from
southern Texas, where it has thus far been recorded from May
to October ; there are many records from Mexico ; also from
Central America, and in South America specimens from Co- '
lombia, Venezuela, Trinidad, Tobago Island recorded in Feb-
ruary and March; British Guiana in September (Dr. Beebe and
John Tee-Van) ; Ecuador; Brazil in September, December, Jan-
uary, May; Peru in March, June, September, October; Bolivia
in February; Paraguay in December and January. Specimens
from Argentina are dated November and December, and it doubt-
less occurs at other dates.

In his: “Synopsis of the Cicadidas of Ecuador,” 1925, Dr.
Frederic W. Goding states regarding the Genus Quesada that:
“One species has been recognized in Ecuador, which is greenish
yellow, with a fuscous spot on the bases of the second and third
apical cells of tegmina ; it is one of our largest species. ’ ’

Dr. Kenneth J. Haywood, Chief of the Department of Ento-
mology, Tucuman, Argentina, has informed me that Quesada

Sept., 1944]

Davis: Cicadas

215

gigas, is, speaking generally, distributed over Argentina north
of a line drawn between Buenos Aires and Mendoza. So far
there are no available records for Chile or Uruguay. The Doctor
states that this fine cicada is called: “ ‘Chichara grande’ (chi-
chara is a common name for the cicada here), ‘Coyoyo,’ or more
commonly 4 Coyuyo, ’ according to what part of this vast country
you find yourself in. ’ ’

Considering the extended distribution of the species, the speci-
mens from the various parts of its wide range are surprisingly
alike, but sometimes differ individually in color. Those from
Brazil, Argentina, etc., are often quite large, with abdomen notice-
ably broad in the males, but Texas specimens may also differ con-
siderably in size.

There is a colored figure of the insect under the name of Tym-
panoterpes gigas in “Biologia Centrali- Americana, ” 1881, with
an interesting account of its song and habits. In : “ Insect Sing-
ers, A Natural History of the Cicadas,” Dr. J. G. Myers devotes
considerable space to an account of this remarkable species, its
song and habits.

In his: “Catalogue of the Cicadidse, ” 1906, Mr. Distant cites a
number of specific names that have been bestowed upon Quesada
gigas, and when more specimens have been collected from the dif-
ferent parts of its remarkable range, and additional field studies
have been made, including time of appearance, it may be dis-
covered that there are some definite geographic races or even an
additional species involved.

In his: “Preliminary Survey of the Cicadidse of the United
States, Antilles and Mexico,” 1892, Uhler stated concerning
gigas : “ I have examined specimens from various parts of Mexico,
and from Guatemala, Guiana, and Matto Grosso, Brazil. Speci-
mens from Tamaulipas, Mexico, differ in no respect from others
living further South, although the species is a very variable one
especially in the amount and form of black marking on the upper
surface of the body. The region of the Bio Grande of Texas is
the most northern limit of this species, while the northern part of
the Argentine Republic seems to be its most southern habitat.”

In: “Notes Del Museo de la Plata,” Tomo V, Buenos Aires,
1940, Prof. Belindo Adolfo Torres described and figured a dark-

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[Vol. LII

colored female of gig as, but as be could find no structural differ-
ence he considered the variety unworthy of a name.

In Texas Quesada gigas has been recorded from Starr, Hidalgo,
Cameron and Bexar counties by Mr. F. F. Bibby, and no doubt it
will be discovered over a wider range. There are specimens from
Kingsville, Kleberg County, in the collection of Cornell Uni-
versity.

Mr. H. B. Parks, Director of the State Agricultural Research
Laboratory near San Antonio, Bexar County records gigas as a
yearly visitor observed since 1934. The “Popcorn Whistlers”
occur in the live-oaks about the Laboratory, usually in July and
August, and he heard them singing in the evening and sometimes
in the early morning in 1941. He has found dead specimens
beneath the trees. He also reports their abundance in the coastal
city of Corpus Christi, Nueces County, in August, 1940. The
little boys in town found great sport in climbing trees and catch-
ing the cicadas. They had also been observed and collected in
1938 at Corpus Christi, and I received specimens from both Mr.
Parks and Mr. Emmett S. Claunch, Jr., who reported that they :
“whistle instead of buzz — that is they sound as though they
whistle.”

Many collectors have found gigas about Brownsville, Cameron
County, and Dr. James A. G. Rehn, of the Philadelphia Academy,
likened its song to the shrill tin whistle of a peanut roaster.

Dr. Raymond H. Beamer and associates from the University
of Kansas collected in Bee County and Hidalgo County in July,
1928, and in sending specimens the Doctor wrote that they had
70 more if I cared to see them. This, as well as some of the other
facts are mentioned to show what a highly successful species
gigas really is, both in numbers as well as in wide distribution,
for in some part of its range from north to south, a male gigas
is in song every month in the year.

Mr. Paul C. Avery of Mission, Hidalgo County, near the Rio
Grande, has sent me a great many gigas, which sometimes occurs
very plentifully along the river where the soil is more or less
damp. He has found them often on Mesquite, and describes the
song as: “Very loud, continuous and shrill. The loudest of any
species found,” at Mission. He collected many in 1935, and in

Sept., 1944]

Davis : Cicadas

217

1936 sent me as a sample 400 specimens — 247 males and 153
females — collected in July of that year. He observed the first
gigas on June 13, and in his letter of July 5, 1936, stated that
they sang both early and late, and often were heard singing after
dark. He heard the last one on September 21 in 1936. Mr.
Avery has also observed the Cicada killer, Sphecius, with a gigas ,
which : ‘ ‘ sure was crying loud and mournfully. ’ ’ This species like
many other cicadas, is subject to a fungus disease, and a number
of specimens have been received with the terminal segments of the
abdomen missing, as often happens when the Seventeen-year
Cicada is attacked by fungus.

It will be seen from the foregoing, that in Quesada gigas the
United States includes in its fauna one of the most remarkable
of the known cicadas, which species is sure to attract more and
more attention in the years to come.

Cornuplura, new genus.

In the Journal of the New York Entomological Society for June,
1936, Tibicen curvispinosa was described and figured as a remarkable cicada
from western Mexico. The two conspicuous upturned spines in the male pro-
truding backward from segment IX, were described and figured, and it was
stated that : 1 ‘ the uncus in curvispinosa is deeply cleft with the resulting two
claw-like extremities long and curved inward. ’ ’ These characters are also
noted by Smith and Grossbeck in : “ Studies in Certain Cicada Species, ’ ’
Entomological News, April, 1907, and shown in their figures 7 and 8 on
plate 3.

Tibicen nigroalbata was also described in the 1936 paper, and its resem-
blance to curvispinosa noted. Only a single female nigroalbata taken by
Prof. E. D. Ball in Arizona was available in 1936.

In the Journal of the New York Entomological Society for June,
1942, these remarkable cicadas are again considered in the light of addi-
tional specimens from Nayarit, Mexico, and Santa Cruz County, Arizona, and
it was suggested that nigroalbata might be a northern race of the southern
curvispinosa.

The general form of the body except for the curved spines in the males of
both curvispinosa and nigroalbata, including the position of the cross-veins
in the fore wings, and the terminal central spine on the last dorsal segment
in the males, is as in the genus Tibicen, but the deeply cleft and curved uncus
is not as found in the other species of that genus native to North America,
nor as in plebeja Scopli, of Europe, the type of the genus. It is suggestive
of the uncus in some species of Diceroprocta, but in that genus the dorsal
segment in the male terminates in two lateral lobes and the venation is
different.

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[Vol. LII

It would appear from the above that a new genus should be erected for
these remarkable insects with curvispinosa as a type, to be placed between
Tibicen and Diceroprocta, for which the name Cornuplura is here proposed.
The genus may also include rudis Walker, from Mexico, as a closely related
species.

The student is referred to several of the structural characters illus-
trated in the above-mentioned three papers, as characteristic of the Genus
Cornuplura.

OKANAGANA SYNODICA (SAY), ITS HABITS, DISTRIBUTION,
AND A NEW COLOR FORM

In 1825 Thomas Say described his Cicada synodica, and stated
that: “Dr. James and Mr. Peale observed this species in great
numbers in one locality at the base of the Rocky Mountains but
it did not occur elsewhere.” He described the body as black
above, and also enumerated the extensive testaceous colored lines
and spots that generally give a number of the insects when seen
together a yellow-brown appearance. Say states : “Scutel [meso-
notum] with a lateral marginal line the elevated X and two dorsal
dilated lines testaceous; the dorsal lines are merely emarginate
on the inner side, and do not form the W ; at the tip of each
anterior line of the X is a conspicuous, black impressed puncture,
and behind the X the posterior edge of the scutel [metanotum] is
visible and testaceous : beneath very pale testaceous. . . . Length
to the tip of the hemelytra less than one inch.”

In the Kansas University Science Bulletin, March, 1920, p. 345,
Dr. P. B. Lawson, in: “The Cicadidse of Kansas,” records syn-
odica from the western part of the state only, and describes it
as : “A small black and honey-yellow species, ’ ’ with length of
body 15 to 18.5 millimeters, and expanse of fore-wings 38 to
44 mm.

Mr. Joseph Duncan Putnam in his : “Remarks on the Habits
of Several Western Cicadae,” Pro. Davenport Academy of Nat-
ural Sciences, March, 1881, records that: “ Cicada synodica Say,
was quite common on the grassy plains near Denver and Boulder,
in Colorado, in June, 1872. The male makes a tolerably loud
rattling noise.” In June, 1920, the late Dr. Lutz, of the Amer-
ican Museum of Natural History, collected four male synodica
at Medicine Bow, Wyoming, about 6,600 feet, and recorded that :
“The small brown cicada in grass has a continuous note, but
sometimes continues for only a short time.”

Sept., 1944]

Davis: Cicadas

219

In June and July of 1935 a brood of this species appeared
in Colfax County, New Mexico. In the writer’s collection there
are 135 specimens representing this brood, and they are all of the
typical brownish-colored form.

In his account of: ‘ ‘ Characteristics of Certain Western
Cicadas,” Jour. N. Y. Ento. Soc., June, 1940, Dr. John W.
Sugden records that : “A large brood of Okanagana synodica was
observed in Emery County, Utah (June 7, 1928). At first, the
sound was mistaken for the humming of the carburetor of the
car. In the field, so many were buzzing that it was impossible
to locate the position of any individuals. After examining the
low, sage-like bushes, thousands of the insects were found.
Fifteen or twenty could be easily collected on a bush not over
a foot high. The note similar to the typical Okanagana song, was
not very loud, but shrill and long continued and what the indi-
vidual lacked in volume was made up for by the large numbers.
Very few would fly if disturbed, but would remain on the bushes
and could be collected by hand. Others would become quiet when
disturbed and fall to the ground, where their color closely re-
sembled the buff-colored soil. Many were mating. The exuvia
were on the ground or attached to the stems. ’ ’

The known range of synodica has been greatly extended in
recent years, and specimens have been examined from Alberta,
Montana, North and South Dakota, Wyoming, Nebraska, western
Kansas, Colorado, Utah, western Texas, New Mexico and Arizona.
It should also be found in Oklahoma as some of the known locali-
ties are close to the state line. About 600 specimens have been ex-
amined, and it has been observed as the specimens accumulated
that the broods appearing in Arizona and New Mexico were some-
times composed of darker-colored individuals than the broods
occurring in Colorado and other more eastern localities.

On May 26, 1941, Mr. Frank H. Parker collected a great many
almost entirely black-bodied individuals at Holbrook, Navajo
County, eastern Arizona, and wrote as follows concerning them :
“The Holbrook series was taken on a large, slightly rolling mesa
covered chiefly with bunch grass, Gutierrezia, and a low (1 foot
or less) shrub somewhat resembling Fairies Feather Duster,
among which was to be found an occasional diminutive Opuntia.

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[Vol. LII

The capacity of my cyanide jars, and time, were the only factors
preventing the capture of many thousands of this species. ’ ’ Mr.
Parker sent 67 males and 33 females from this brood.

In the writer’s collection there are also two females of the dark
form collected at Holbrook, May 22, 1934, and a male and female
of the same form from the White Mountains, Arizona, July, 1935.
Broods of both the light and dark forms occur in New Mexico,
and both dark and light colored specimens have been received
from Utah, collected by Dr. Sugden, June 7, 1928, in Emery
County.

It will be noted that broods of the dark, or almost wholly
black individuals, generally occur in the area drained by the
Colorado, while those of the lighter-colored form described by
Say, are on the more eastern watershed, or streams tributary to
the Rio Grande, Missouri, etc.

Okanagana synodica (Say) variety nigra, new variety (Plate VIII, tig. 3).
Type male and allotype female from Holbrook, Arizona, May 26, 1941
(Frank H. Parker). Davis collection.

In this dark or melanistic form of synodica, the testaceous markings as
described by Say are absent or very much reduced. The head is entirely
black save for two pale spots one above each antenna, and the ruby-colored
ocelli. The pronotum is narrowly edged all around with orange, and has a
short, median pale line extending to the anterior margin. The mesonotum
has a very small orange spot at the base of each fore wing ; the posterior
margin, including the X, pale, and the two torch-shaped orange marks, ex-
tending forward from the anterior limbs of the X generally lack the often
conspicuous tooth-like mark or sinuation on the inner side found in typical
synodica. The metanotum is pale and the abdominal segments are black or
very narrowly margined posteriorly with orange. In the female the seg-
ments at the end of the body are more broadly margined with orange. The
upper surface of the uncus of the male is black, and the valve pale. The
venation is darker than in typical synodica.

OKANAGANA PALLIDULA DAVIS ; ADDITIONAL NOTES
AND OBSERVATIONS

(Plate VIII, Fig. 4)

In the Journal of the New York Entomological Society for
September, 1938, there is an account of this species, its distribu-
tion and color forms, and the variety nigra is described and re-
corded from Yolo and Kern counties, California. Pallidula can
be confused with some of the forms of Okanagana vanduzeei

Sept., 1944]

Davis: Cicadas

221

which, however, are usually conspicuously hairy on the head, ancl
often on the pronotum, as well as on the under side of the ab-
domen. Also pallidula is duller and does not present the shin-
ing appearance usual in vanduzeei and its forms. While there
are dark specimens of pallidula there are also green ones, which
apparently do not occur in vanduzeei and its varieties eonsoibrina,
etc.

Okanagana pallidula has thus far been examined from the
great Central Valley of California, and from the following coun-
ties: Sutter, Yuba, Yolo, Sacramento, Contra Costa, Merced,
Madera, Fresno, Tulare, Kings, Kern and San Luis Obisbo. In
some years it occurs in great numbers.

As reported on page 308 of the September, 1938, paper referred
to above, this small cicada may be found singing from its hole in
the ground. In June, 1941, Mr. and Mrs. J. N. Knull of the
Ohio State University, while in the Santa Maria River Valley,
California also found specimens of pallidula singing from their
burrows where it was believed they had undergone their last trans-
formation. If the males ultimately found mates, they prob-
ably in due time left the burrows, or perhaps the females flew to
them as has been observed in other species. This is a matter for
future observation and record. Dr. Charles D. Michener of the
American Museum of Natural History, while looking over my
collection of cicadas in 1942, assured me that he had discovered
males of pallidula singing from their one-time burrows, thus
adding to the observations made by others.

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Journal New York Entomological Society

[Yol. lii

Plate YIII

Figure 1. Quesada gigas (Olivier). Male from Texas.
Figure 2. Quesada gigas (Olivier). Female from Argentina.
Figure 3. Olcanagana synodica (Say) variety nigra. Type.
Figure 4. Olcanagana pallidula Davis.

(Jour. N. Y. Ent. Soc.), Vol. LIT

(Plate YIII)

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate IX)

EPHRAIM. PORTER FELT

Sept., 1944]

Bromley: Felt

223

EPHRAIM PORTER FELT— 1868-1943

The New York Entomological Society as well as the entire
scientific world has lost one of its most respected and outstanding
members in the passing of Dr. E. Porter Felt. He was a leader
in the field of entomology and was recognized as such inter-
nationally. He was widely known as a scientist, writer and lec-
turer. As State Entomologist of New York for thirty years his
valuable publications were looked upon as standard for this type
of work. As Director of the Bartlett Tree Research Laboratories
from 1928 until his death, he became a world authority on the
care of shade trees. Real leaders are few in any walk of life.
Dr. Felt was one of the few but his influence was valued by many.

Ephraim Porter Felt was born at Salem, Massachusetts, Janu-
ary 7, 1868, a son of Charles Wilson Felt and Martha Seeth
(Ropes) Felt. His background was like one of many similar old
New England families. As a youth he had decided upon the
ministry as a career but became interested in insect problems
while attending college and this became the basis of his life work.
He received the degree of B.Sc. from the Massachusetts Agricul-
tural College in 1891 and also a degree of B.Sc. from Boston Uni-
versity the same year. The degree of D.Sc. was bestowed upon
him by Cornell University in 1894. After teaching Natural Sci-
ence at The Clinton Liberal Institute, Fort Plain, New York,
from 1893-1895, he became Assistant to the State Entomologist
at Albany. Upon Dr. Lintner’s death, he became State Ento-
mologist of New York, which post he held for thirty years, until
1928 when he came to the Bartlett Tree Research Laboratories as
Director and Chief Entomologist.

In the field of insect taxonomy he achieved international fame
for his careful and extensive studies on the . gall midges — the
Cecidomyidse or Itonididas as he preferred to call them. In 1913
he gave the annual public address on Gall Insects before the Ento-
mological Society of America at Atlanta, Georgia. In 1914 he
was elected a member of the American National Committee on
Nomenclature of the Second International Congress. I have
learned from Professor C. P. Alexander of Massachusetts State

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Journal New York Entomological Society

[Vol. LII

College that Dr. Felt had reported to him in 1942 that he had
described 1,060 new species of gall midges and plant mites. With
the late D. B. Young, his assistant at Albany, he had described
a number of mosquitoes which are indicated in the literature
under the joint authorship of Felt and Young.

Dr. Felt became a corresponding member of the New York
Entomological Society on October 20, 1900, and an active member
on February 5, 1907. After coming to Stamford in 1928 he fre-
quently presented papers at the New York Entomological Society
meetings which included subjects of a varied nature from gall
midges to poetry concerning insects, as well as numerous valuable
contributions on shade tree insects.

Dr. Felt’s first entomological work was in Massachusetts in the
early days of the gypsy moth invasion. His experience with this
destructive forest and shade tree pest led him in 1924 in his work
for the New York State Conservation Commission to propose a
barrier zone two hundred miles in length to stem the spread of
the gypsy moth into New York State from New England.

Dr. Felt was a pioneer in airplane collection of insects and
made extensive studies on windborne insects. He made studies
on the wind drift of insects on the top of the State Education
Building, Albany, N. Y., the Empire State Building in New York
City, as well as on other tall city buildings and his work on wind-
borne insects has been very important in ascertaining the direc-
tion of spread of the Dutch elm disease which is carried to a large
extent by the European elm bark beetle.

In the late 90 ’s and early 1900 ’s, Dr. Felt studied the spread
of many diseases by the common house fly and by mosquitoes and
made important contributions to this phase of entomology. In the
February, 1944, issue of the National Geographic there was an
article “Saboteur Mosquitoes” by Mr. Stage, Senior Entomolo-
gist of the United States Bureau of Entomology and Plant
Quarantine, in which is recorded some of the work done on mos-
quitoes years ago under the direction of Dr. Felt.

Dr. Felt was very much interested in and very capable of doing
editorial work and was editor of the Journal of Economic Ento-
mology since its start in 1908. At one time he was editor of the
National Shade Tree Conference Reports.

Sept., 1944]

Bromley: Felt

225

While State Entomologist of New York he published 25 official
reports covering the activities of that office and many bulletins,
which were published by the New York State Museum, as well as
more than 700 timely papers and articles which included a wide
variety of entomological work comprising general entomology as
well as popular articles of an informative nature for the general
public. He was a frequent contributor to entomological journals
and scientific publications, as well as magazines and newspapers.
He found time to give numerous addresses, to publish several
popular books and to speak over the radio on different phases of
entomological and shade tree activities. His published reports
and bulletins were models of systematic arrangement and clarity
of expression.

During his long and useful career, Dr. Felt had been a collabo-
rator of the United States Bureau of Entomology, the entomologi-
cal editor of the Country Gentleman from 1898 to 1911 and a
member of the Supervisory Board of the American Year Book.
He was a past president of the American Association of Economic
Entomologists, a past president of the National Shade Tree Con-
ference, a fellow of the Entomological Society of America, a life
member of the American Entomological Society, a member of the
New York and Washington Entomological Societies and of Sigma
Xi and Phi Kappa Phi. Among the honors accorded him were a
gold, and three silver medals at the Pan-American Exposition,
Buffalo, N. Y., in 1901. He was a member of the Fraternity of
Alpha Sigma Phi.

For the last twelve years, he was the author of a weekly syndi-
cated newspaper article entitled “ Talks on Trees” which was used
by nearly three hundred newspapers throughout the country.
He was the author of a number of well-known books on insects
and trees, his two-volume work “Insects Affecting Park and
Woodland Trees” appearing in 1906 and “Manual of Tree and
Shrub Insects” in 1923. He was co-author with Dr. W. H.
Kankin of “Insects and Diseases of Ornamental Trees and
Shrubs” published in 1932. His “Plant Galls and Gall Makers”
appeared in 1940.

Three outstanding books on shade trees were published more
recently — 1 1 Our Shade Trees, ’ ’ in 1938, a second edition of which

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Journal New York Entomological Society

[Vol. LII

appeared in 1942; “Pruning Trees and Shrubs,” in 1941, and
“Shelter Trees in War and Peace,” in 1943.

On December 8, 1943, Dr. Pelt attended the national meetings
of the American Association of Economic Entomologists held in
Columbus, Ohio. On his way back he stopped at Utica, N. Y.,
to visit his son. He was back at his office on Monday, December
13. He seemed none the worse for the rigorous trip and was in
excellent spirits. On Tuesday morning, December 14, he told his
secretary that the night before he doubted whether he would be
in that day. He said he felt as though he was coming down with
the flu the night before but felt all right that morning. With a
twinkle in his eye, he said that he had had such symptoms before
and that nothing came of them. “You see,” he said, “I am still
pretty tough.” These were the last words that Miss Burns, his
secretary, heard him speak. I talked with him a few minutes
later when he was going out to the Ferguson Library to look up
some references and he seemed to be all right then. Later he
came back to the building and died very suddenly of a heart
attack.

Surviving him, in his immediate family, are three daughters
and a son, a sister and two brothers, and thirteen grandchildren.

A few days after Dr. Felt had passed away, I received a letter
from Dr. Peairs, present editor of the Journal of Economic Ento-
mology, recounting his conversation with Dr. Felt during the
Columbus meetings. I would like to quote that part of Dr.
Peair’s letter.

“I cannot refrain from telling you a little episode in Columbus ;
I had a chat with Dr. Felt and he brought up a discussion of some
of the members who had recently died ; I casually mentioned the
fact that four ex-presidents had died in the last few months. Dr.
Felt, with his faint smile, remarked, not seriously at all, ‘I am
hereby resigning my position as an ex-president of the Associa-
tion, effective at once. ’ I am sure he thought of it as nothing but
a little joke, as I did. But somehow, I almost wish the subject
had not come up. ’ ’

Dr. Felt was a great scientist, a splendid gentleman and an
invaluable friend. Honest, sincere and upright, he moved
through life with a tranquil demeanor and faith in his fellow

Sept., 1944]

Bromley: Felt

227

men. Trivialities neither upset him nor deterred him from any
worthy purpose or goal.

His prime thoughts were for the younger, newer men. All were
stimulated by his great personality. He aided their efforts with
cheer and lent a helping hand no matter how adverse the diffi-
culty or how uncertain the going.

I first met Dr. Felt in 1920. I have known hinqas a friend, as
a counsellor, as a superior and as a leader. Since 1929, it was
my privilege and honor to work with him side by side on impor-
tant research and I have never ceased to marvel at his deft ability
to size up and conquer a problem and at his unruffled composure
in the face of trying obstacles.

Of him I can use no greater expression than to say that he was
an all around great American. His flexibility of outlook was
manifested by the ability to direct itself in a flash from the deep
abstruse contemplation of science to the broad generalities of
national life. He was intensely interested in the world about him.
His human ties were broad and comprehensive. His love for
children was apparent to many.

He was an avid reader, and enjoyed the radio from the political
speeches of the great to the humor of Charlie McCarthy. He had
a deep and abiding sense of humor and was quick and accurate
at repartee. A remark was once made on his winged collar, so
characteristic of his dress. Dr. Felt immediately replied 1 1 What
could be more fitting to my profession?” pointing to the two-
winged insects he was studying.

He was faithful to the minute to his commitments. Many were
the occasions when he accepted an invitation to talk at some
humble, out-of-the-way garden club or other meeting. If he said
he would be there, he was always present. This custom alone
stamped him as a man of his word in the eyes of many of his great
circle of friends.

His appearance as well as his character was left unchanged by
the surging tide of years and he was until the last the same stead-
fast, striking figure. His carefully trimmed, white hair and Van
Dyke beard, his calm, upright bearing, his impeccable dress, his
gentle but firm voice, all reflected the imprint of his great
character.

228

Journal New York Entomological Society

[Vol. LIT

How frequently and forcefully the realization has come to me
during the past two months that we will never again experience
together the exhilaration of solving an important problem, or of
unravelling one of nature’s intricacies again; that no more will
we talk over together the problems of the present or the plans of
the future ; that no more will I have his steadfast character upon
which to lean or his guiding hand to point the way !

While his passing has been such a personal loss to me, how
much greater has been the loss to the profession of entomology
and to the entire world of science. His contributions will loom
in importance during the passing years even as the memory of his
personality may dim and fade.

The accompanying bibliography, transcribed from the card
index by his secretary, Miss Beatrice Burns, gives his publications
in chronological order. It is an inescapable conclusion that such
a bibliography cannot be complete. Dr. Felt was accredited with
having written more than seven hundred important and timely
papers and articles which include a wide variety of entomological
works.

The following may be considered his outstanding writings :

BIBLIOGRAPHY OF DR. E. P. FELT

1893. Nomophila noctuella. Can. Ent. 25 : 129.

1894. On certain grass-eating insects. Cornell Agric. Exp. Sta. Bull. 64.
1896. The scorpion flies. (10th Bep’t) N. Y. State Ent. for 1894.

1901. Suggestions toward greater uniformity in nursery inspection laws and

rulings. Cony, of the Assoc, of Amer. Agr. Colleges and Ex-
periment Stations, Bui. No. 99.

A 1902. N. Y. State Mus. Bui. 53 (17th Rep’t), 705-30, 741-44.

1902. Crude petroleum as an insecticide. Twenty-Third Meet. Soc. Prom.

Agr. Sc. p. 1.

1903. Literature of American economic entomology. Bureau of Entomology

Bui. 40, new series, 7—22.

1904. (With L. H. Joutel) Monograph of the genus Saperda. N. Y. State

Mus. Bull. 74, 1-80.

1904. Importance of isolated rearings from culicid larvge. Ent. Soc. Wash.
Proc. VI, Vol. XX, 312-13.

1904. The mosquitoes or culicidse of New York State. N. Y. State Mus.
Bull. 79, 241-400, 113 fig., 57 pi.

1904. The cause and control of insect depredations. Twenty-Fifth Ann.
Meet. Soc. for Promotion Agr. Sc., p. 73.

Sept., 1944]

Bromley: Felt

229

X 1905.
1906.

1906.

X 1906.

1906.

1907.

1907.
& 1907.

X 1908.

1908.
1908.

? 1908.
1908.
1908.

1908.

1910.

1911.
1911.

1911.
1911.
1911.
1911.
1911.
1911.
XL 1911.

Cecidomyia hirtipes. N. Y. State Mus. Bull. 97 (Rep’t 1904), 410-11.
Diversities among New York mosquitoes. Proc. 2nd Anti-Mosquito
Conv. New York City, 1904.

Mosquito control. N. Y. State Mus. Bull. 104 (21st Rep’t), N. Y.
State Ent., 1905.

Studies in Cecidomyiidae. N. Y. State Mus. Bull. 104 (21st Rep ’t,
1905), 116-32.

Plant galls & gall makers. N. Y. State Mus. Memoir 8, 2: 615-48.
Insects affecting park & woodland trees. N. Y. State Mus. Mem. 8,
vol. 1, 1-332.

Hort. diseases and pests. N. Y. State Ed. Dept. Rev. of Legislation,
119-22.

New species of Cecidomyiidae. N. Y. State Mus. Bull. 110, 97-165.

Separate, 1-53. Lingnan Science Jour., Yol. 7., 469.

Gall gnats or Cecidomyiidae. Can. Ent., 39: 143-44.

Cecidomyiidae : A statement. Can. Ent., 39 : 197-98.

Cecidomyia acarivora n. sp. Ent. News, 17 : 242.

New species of Cecidomyiidae II, p. 1-23. Same in N. Y. State Mus.
Bull. 124, 286-304,

Cecidomyia johnsonii Sling. Jour. Econ. Ent., 1: 243.

Contarinia gossypii n. sp. Ent. News, 19 : 210-11.

Insect control in its larger aspects. Fruit Growers Proceedings, p. 1.
Some problems in nomenclature. Ann. Ent. Soc. of America, p. 102.
Observations on the genus Contarinia . Jour. Econ. Ent., 1: 225-28.
Observations on the biology and food habits of the Cecidomyiidae.
Jour. Econ. Ent., 1: 18-21.

Studies in Cecidomyiidae II. N. Y. State Mus. Bull. 124 (23d Rep’t),
286-304.

New species of Cecidomyiidae II. N. Y. State Mus. Bull. 124, 307-510.
Schizomyia ipomoece n. sp. Ent. News, 21: 160-61.

Gall midges of Aster, Carya, Quercus and Salix. Jour. Econ. Ent.,
3: 347-56.

West Indian Cecidomyiidae. Ent. News, 21: 268-70.

Two new Cecidomyiidae. Ent. News, 21: 10-12.

Observations on the house fly. Jour. Econ. Ent., 3: 24-26.

A generic synopsis of the Itonidae. Jour. N. Y. Ent. Soc., 19: 31-62.
Summary of food habits of American gall midges. Ann. Ent. Soc.
America, 4: 55-62.

Endaphis Kieff., in the Americas (Dipt.). Ent. News, 22: 128-29.
Endaphis hirta n. sp. (Dipt.). Ent. News, 22: 224.

Two new gall midges. Can. Ent., 43: 194-96, June.

Four new gall midges (Dipt.). Ent. News, 22: 301-5.

Hosts and galls of American gall midges. Jour. Econ. Ent., 4: 451-75.
New species of gall midges. Jour. Econ. Ent., 4: 476—84; 546-59.
Miastor americana Felt, an account of pedogenesis, N. Y. State Mus.
Bull. 147 (26th Rep’t), 82-104.

230

Journal New York Entomological Society

[Yol. lii

1911. A new species of Lasioptera with observations on certain homologies.
Psyche, 18: 84-86.

¥• 1911. New Indian gall midges. Mem. Dept. Agr. India Ent. Ser. VII, No.
4, 23-28.

1911. A new Lestodiplosis. Ent. News, 22 : 10-11.

1911. Two new gall midges (Dipt.) Ent. News, 22: 109-11.

1911. Three new gall midges. Jour. N. Y. Ent. Soc., 19: 190-3.

1911. Rhopalomyia grossularicB. Jour. Econ. Ent., 4: 347.

1912. New W. Indian gall midges (Dipt.). Ent. News, 23: 173-7.

1912. Observations on Uleella Rubs. (Dipt.). Ent. News, 23: 353-54.

1912. Lasiopteryx manihot n. sp. Can. Ent. 44: 144.

1912. Studies in Itonididse. Jour. N. Y. Ent. Soc., 20: 236-48.

1912. Observations on the identity of the wheat midge. Jour. Econ. Ent.,
5: 286-89.

1912. The gall midge fauna of western North America. Pomona Coll. Jour.
Ent., 4: 753-57.

1912. The identity of the better known midge galls. Ottawa Nat., 25:
65-67, 181-88; separate, 1-11.

1912. Biology of Miastor and Oligarces. Science, 33 : 278-80.

1912. Diarthronomyia calif ornica n. sp. Pomona Coll. Jour, of Ent., 4: 752.
1912. Petroleum & petroleum products as insecticides. N. Y. State Ed.
Dept., Sc. Div. 2 p.

1912. Itonida inopis. Jour. Econ. Ent., 5: 368—69.

1912. Control of insect pests in institutions. Jour, of Home Econ., Yol.
IY, No. 1, 16-26.

1912. New Itonidse. Jour. N. Y. Ent. Soc., 20: 102-7.

1912. New gall midges or Itonidse (Dipt.). Jour. N. Y. Ent. Soc., 20: 102-7.
1912. New gall midges or Itonidse (Dipt.). Jour. N. Y. Ent. Soc., 20:
146-56.

1912. Arthrocnodax occidentalis. Jour. Econ. Ent., 5: 402.

1912. Observations on the identity of the wheat midge. Jour. Econ. Ent. 5:

286-9.

1913. Cystodiplosis eugenice n. sp. Ent. News, 24: 175-76.

1913. Adaptation in the gall midges. Can. Ent. 45: 371-79; also Ent. Soc.

Ontario (44 Rep’t), 1913, 1914, 76-82.

1913. The gall midge fauna of New England. Psyche, 20: 133-47.

1913. A study of gall midges. N. Y. State Mus. Bull. 165 (28th Rep’t),
State Ent., p. 127-227.

1913. Three new gall midges (Dipt.). Can. Ent. 45: 304-8.

7C 1913. A study of gall midges. N. Y. State Mus. Bull. 175 (29th Rep’t),
79-214.

1913. Gall midges in an aquatic or semiaquatic environment. Jour. N. Y.
Ent. Soc., 21: 62-63.

1913. Table of hickory leaf midge galls. Bui. Bklyn. Ent. Soc., 8 : 98-99.
1913. Two new Canadian gall midges. Can. Ent., 40 : 417-18.

1913. Didactylomyia capitata n. sp. Psyche, 20: 174.

Sept., 1944]

Bromley: Felt

231

1913. Descriptions of gall midges (Dipt.). Jour. N. Y. Ent. Soc., 21:
213-19.

1913. Itonida anthici n. sp. Jour. Econ. Ent., 6: 278-79.

1913. Gouty pine midge. Jour. Econ. Ent., 6: 278-79.
x 1913. The goldenrod and its gall flies. Guide to Nature, 6: 149-51.

1913. Arthrocnodax Carolina. Jour. Econ. Ent., 6: 488-89.

1914. Cecidomyiidae by J. J. Kieffer. Fasc. 152 in Genera Insectorum.

Ent. News, 25: 185-88.

1914. Additions to the gall midge fauna of New England. Psyche, 20:
109-14.

1914. Acaroletes pseudococci. Jour. Econ! Ent., 7: 148-49.

1914. Diadiplosis coccidivora. Entomologist, 47 : 86.

1914. Aplonyx sarcobati n. sp. Pomona Jour. Ent. & Zool., 6: 93—94.
1914. Gall midges as forest insects. Ottawa Nat., 28: 76-79.

1914. List of zoophagous Itonididse. Jour. Econ. Ent., 7 : 458-59.

X 1914. Cactus midge, Itonida opuntice. Prickly Pear Trav. Comm. Rep’t,
77-78.

¥ 1914. Notes on forest insects. Jour. Econ. Ent., 7: 373-75.

1914. Additions to the gall midge fauna of New England. Psyche, 20:
109-14.

1914. Descriptions of gall midges. Jour. N. Y. Ent. Soc., 22: 124-34.

1914. New gall midges (Itonidae). Insec. Inscit. Menstr., 2: 117-23.

1914. Hormomyia bulla n. sp. Can. Ent., 46: 286-87.

1914. Conical grape gall Cecidomyia viticola. Jour. Econ. Ent., 7 : 339.

1914. Arthrocnodax constricta. Jour. Econ. Ent., 7: 481.

1915. New genera and species of gall midges. U. S. Nat. Mus. Proc., 48:

195-211.

1915. New South American gall midges. Psyche, 22: No. 5.

1915. The gall midges of the pine. Bull. Bklyn. Ent. Soc., 10: 74-76.

X 1915. A study of gall midges II. N. Y. State Mus. Bull. 175 (29th Rep’t),
79-213.

i. 1915. A study of gall midges III. N. Y. State Mus. Bull. 180 (30th Rep’t),
127-289.

1915. New Asian gall midges. Jour. N. Y. Ent. Soc., 23: 73-84.

1915. Fumigation for the box leaf miner. Jour. Econ. Ent., 8: 94-95.

1915. Mycodiplosis macregori n. sp. Jour. Econ. Ent., 8: 149.

* 1915. A new chrysanthemum pest. Am. Florist, 44 : 162.

1915. New North American gall midges. Can. Ent. 47 : 236-42.

X 1915. New gall midges. Jour. Econ. Ent., 8: 405-9.

■K 1916. A study of gall midges III. N. Y. State Mus. Bull. 180 (30th Rept’t),
127-288.

A 1916. A study of gall midges IV-. N. Y. State Mus. Bull. 186 (31st Rep’t),
101-73.

1916. New western gall midges. Jour. N. Y. Ent. Soc., 24: 175-96.

1916. Gall midges of certain Chenopodiaceae (Dipt.). Ent. News, 27: 201-3.

1916. New North American gall midges. Ent. News, 17 : 412-17.

232

Journal New York Entomological Society

[Vol. LII

1916.
H 1916.

1916.
■V 1916.
* 1916.

1917.
1917.
1917.
1917.

^ 1917.

1917.
X 1918.

1918.

X1918.
f 1918.

A 1918.
-V 1918.

/ 1918.

1919.
X 1919.
X 1919.

-f 1919.

1919.

1920.
4. / 1920.

1920.

1920.
* 1921.

x( 1921.

X 1921.

New Indian gall midges. Can. Ent., 48 : 400-6.

Distribution of gall midges. Natl. Acad. Sci. Proc., 3 : 349-54.

New gall midges. Can. Ent., 48 : 29-34.

Lasioptera fructuaria. Maine Agr. Expt. Sta. Bull. 244, 268-69.

American insect galls. Ottawa Nat. 30 : 37-39.

Indian gall midges (Cecidomyiidse, Dipt.). Ent. News, 28: 73-76.

Entomological research and utility. Scientific Monthly, p. 551.

New gall midges. Jour. N. Y. Ent. Soc., 25: 193-96.

Distribution of gall midges. Natl. Acad. Sci. Proc., 3: 349-54.

Little known gall midges of certain composites. Ottawa Nat., 31:
13-14.

Asphondylia websteri n. sp. Jour. Econ. Ent., 10: 562.

A study of gall midges Y. N. Y. State Mus. Bull. 198 (32d Rep’t),
101-252.

Notes and descriptions of Itonididse in the collections of the American
Mus. of Natural History. Am. Mus. Nat. Hist., 38 (Oct. 6) :
179-82.

Gall insects and their relation to plants. Sci. Mon., 6 : 509-25.

New Philippine gall midges, with a key to the Itonididse. Phil. Jour.
Sci., 13: 281-325.

Key to American insect galls. N. Y. State Mus. Bull. 200, 1-310.

A study of gall midges YI. N. Y. State Mus. Bull. 202 (33d Rep’t),
76-205.

New gall midges. Jour. Econ. Ent., 11: 380-84.

Five non-gall making midges. Ent. News, 30 : 219-23.

New Philippine gall midges. Jour. Sci., 14: 287—94.

The plant galls collected by the Canadian Arctic Expedition 1913-18.
Kept. Can. Arctic Exped., Y. 3.

Insect galls & gall insects. Ottawa Nat., 32: 127-31.

The European corn borer. Cornell Ext. Bull. 31, 35-48.

The European corn borer problem. Jour. Econ. Ent., 13: 59-74.

Memoirs Department of Agriculture in India. Yol. YII — No. 1, for
Imperial Department of Agr. in India.

Four new African gall midges. Annals Natl. Mus., Yol. IY (part 2),
p. 491-96.

New Indian gall midges. Mem. Dept. Agr. India. Ent. Ser. YII,
No. 1, p. 1-11.

New Philippine gall midges. Phil. Jour. Sci., 17 : 231-34.

Three new sub-tropical gall midges (Itonidae Dipt.). Ent. News, 22:
141-43.

Observations on J ohnsonomyia Felt with a description of a new spe-
cies. Can. Ent., 53 : 96.

New species of reared gall midges (Itonididse). Jour. N. Y. Ent.
Soc., 29: No. 2, 165.

A study of gall midges YII. N. Y. State Mus. Bull. (34th Rep’t),
81-241.

Sept., 1944]

Bromley: Felt

233

1 1921.

\ 1921.
/ 1921.
X 1921.

X 1921.

1921.

1922.
1922.

*1922.
\ 1922.

1924.

1925.

V 1925‘

1925.

* 1926.

* 1926.
^ 1926.

1927.
X 1927.

1928.

1929.

1929.
#1929.

1930.
1930.

1930.

1931.

Adaptations among insects of field and forest. Scientific Monthly,
13: No. 2, 165.

Lasioptera apocyni, new species (Diptera). Can. Ent., 53: 149.

NeAv Javanese gall midges. Extrait De Treubia, Yol. II, p. 89-92.
The number of antennal segments in gall midges and a new species.
Bull. Bklyn. Ent. Soc., 16: 93-95.

Indian grass gall midges. Mem. Dept. Agr. India, Pusa Ent. Ser.
VI, No. 3, p. 15-22.

Javanese gall midges, Treubia, 1: 139-51.

A new Javanese gall midge. Treubia, 1: 270-71.

A new Diadiplosis. Zoologica, Vol. Ill, No. 8.

The possibilities of exterminating insects. Scientific Monthly, 15:
No. 1, 35-41.

A new and remarkable fig midge. The Ent. (Summer Number), p.
5-6.

New cecidomyiid parasite of the whitefly. U. S. Natl. Mus. Proc.,
61: 1-2.

Manual of tree & shrub insects. The Macmillan Co., N. Y., 382 p.
School guide to insects and about insects. Univ. of State of N. Y.
Press.

Key to gall midges. (A Besume of Studies I— VII, Itonididae). N. Y.

State Mus. Bull. No. 257, 1-239.

Dispersal of butterflies and other insects. Nature, p. 1.

Insects and human welfare. Scientific Monthly, 21: 649-53.

A new spruce gall midge (Itonididae). Can. Ent., 58: 229.

New non-gall making Itonididae (Dipt.). Can. Ent., 58: 265-66.

New species of Indian gall midges (Itonididae). Memoirs of the Dept,
of Agr. in India, 9 : 241-45.

Four new Indian gall midges. Mem. Dept. Agr. India, 10 : 1-4.

New species of East Indian gall midges. Treubia, 9: 385-89.

A new African gall midge. Jour. N. Y. Ent. Soc., 36: (2), 123-24.
The need for investigations on shade tree insects. Jour. Econ. Ent.,
22: No. 2, 417.

Hickory leaf gall midges Caryomyia species. Jour. Econ. Ent., 22:
422.

Gall midges or gall gnats of the Orient (Itonidae or Cecidomyidae)
Lingnan Sc. Jour., Vol. 7, 413-74.

The Norway maple Nepticula (Lepidoptera) . Proc. Ent. Soc. Wash.,
32: 8, 146-9.

Economic importance of shade tree insects. Jour. Econ. Ent., 23:
1, 109-13.

(With S. W. Bromley) Shade tree problems. Proc. of the Ann.

Meeting of the Natl. Shade Tree Conf., p. 13.

Scientific names. Science, 21 : No. 1834, 215-17.

(With S. W. Bromley) Developing resistance or tolerance to insect
attack. Jour. Econ. Ent., 24: 437.

234

Journal New York Entomological Society

[Yol. lii

1931.

1932.

1933.

1934.

1935.

n 1935.

1935.

(^With S. W. Bromley) Insecticide investigations during 1930. Jour.
Econ. Ent., 24: 232.

The effect of certain important insects on trees. Natl. Shade Tree
Conf., 66-73.

(With S. W. Bromley) Observations on shade tree insects. Jour.
Econ. Ent., 24: 157-62.

(With S. W. Bromley) Tests with nicotine activators. Jour. Econ.
Ent., 24: 105.

(With W. Howard Rankin) Insects and diseases of ornamental trees
and shrubs. The Macmillan Co., N. Y., 507 p.

A new citrus cambium miner from Puerto Rico. Jour. Dept, of Agr.

of Puerto Rico, Yol. XYI, No. 2, 117-18.

A new predaceous gall midge for California. Pan-Pacific Ent., 8:
No. 4, 167-68.

A new Japanese gall midge. Bull. Bklyn. Ent. Soc., 27 : No. 2, 124.
Tree insurance. Proc. Ann. Meet. Natl. Shade Tree Conf., p. 15.
Shade tree insects. Proc. Ann. Meet. Natl. Shade Tree Conf., p. 77.
Shade trees threatened by insect pests. Scientific Monthly, 35 : 59-62.
(With S. W. Bromley) Insecticides on shade trees and ornamentals.
Jour. Econ. Ent., 25: 298.

(With S. W. Bromley) Observations on shade tree insects. Jour.
Econ. Ent., 25 : 39.

A hibiscus bud midge new to Hawaii. Proc. Haw. Ent. Soc., VIII,
No. 2, 247-48.

A new enemy of the pineapple mealybug and a list of gall midge
enemies of mealybugs. Jour. N. Y. Ent. Soc., 41: Mar .-June,
187-89.

New gall midges. Bull. Bklyn. Ent. Soc., 29: No. 2, 77-78.
Fundamentals in the control of insect pests. Proc. 10th Natl. Shade
Tree Conf., 37-43.

Classifying symbols for insects. Jour. N. Y. Ent. Soc., 42: 373-92.

A new gall midge on fig. (Dipt.: Itonididse). Ent. News, 45: 131-33.
Shade tree insects in 1933. Jour. Econ. Ent., 27 : 195.

Bark beetles and the Dutch elm disease. Jour. Econ. Ent., 28: No. 1,
231.

(With K. F. Chamberlain) Occurrence of insects at some height in
air, especially on the roofs of high buildings. N. Y. State Mus.
Circ. 17, 70 p.

Trisopsis in the United States (Dipt.: Itonididae or Cecidomyiidae),
Ent. News, 46: 75.

A new melon gall midge. Bull. Bklyn. Ent. Soc., 30: No. 2, 79-80.

A new gall midge. Jour. N. Y. Ent. Soc., 43: 47.

A gall midge on pine cones. Jour. N. Y. Ent. Soc., 43: 48.

Bark beetles and the Dutch elm disease. Jour. Econ. Ent., 28: 231.
The importance of shade tree insects in 1934. Jour. Econ. Ent., 28:
390.

Sept., 1944]

Bromley: Felt

235

1936.

1937.

1938.

1939.

1940.

X 1940.
1940.
1940.

1940.

1940.
’*1941.

1941.
)C 1941.

1941.

New midges on pine and grass. Jour. N. Y. Ent. Soc., 44: 7.

(With S. W. Bromley) Oil effects on shade trees. Jour. Econ. Ent.,
29: (2), 357-60.

Two new cockle burr midges (Dipt.: Cecidomyiidae) . Ent. News, 47:
231.

(With S. W. Bromley) Shade tree insect developments. Jour. Econ.
Ent. 29: 490.

The biology of mayflies, with a systematic account of North American
species. Jour. Econ. Ent., 29: 223.

Forest insects. Jour. Econ. Ent., 29: 1031.

(With S. W. Bromley) Observations on shade tree insects and their
control. Jour. Econ. Ent., 30: (1), 71-75.

Dissemination of insects by air currents. Jour. Econ. Ent., 30: 458.
A note on Lasioptera murtfeldtiana Felt. Jour. N. Y. Ent. Soc.,
46: 44.

(With S. W. Bromley) Shade tree insects and sprays, 1937. Jour.
Econ. Ent., 31: No. 2, 173-6.

Why do insects become pests? Scientific Monthly, 46: 437-40.

A new species of gall midge predaceous on mealybugs. Proc. Haw.
Ent. Soc., No. 1.

Wind drift & dissemination of insects. Excerpt, Can. Ent., 221.

Our shade trees. Orange Judd Publishing Company, N. Y., 187 p.
(With S. W. Bromley) Hurricane of Sept. 21, 1938. Scientific Tree
Topics No. 1, 1939, Yol. 1.

A new juniper midge (Diptera: Cecidomyiidae). Ent. News, 50:
159-60.

A new gall midge on rhododendron. Jour. N. Y. Ent. Soc., 47 : 41-42.
(With S. W. Bromley) New insecticides and spreaders on shade trees.
Jour. Econ. Ent., 33: 244-49.

Rhododendron midge. Scientific Tree Topics 4, Yol. 1, p. 30.
Insecticides and their use. Scientific Tree Topics 4, Yol. 1, p. 26.
European elm bark beetle and Dutch elm disease control. Jour. Econ.
Ent., 33: (3), 556.

Danger to trees in an oversupply of quickly available nitrogen. Scien-
tific Tree Topics 2, Yol. 1, p. 10.

Pine root weevil. Scientific Tree Topics 2, Yol. 1, p. 13.

Factors affecting Dutch elm disease spread. Scientific Tree Topics
3, Vol. 1. p. 18.

Training trees for beauty. Scientific Tree Topics 5, Yol. 1, p. 34.
(With S. W. Bromley) New and unusual shade tree pests. Jour.
Econ. Ent., 34 : (3), 383.

With S. W. Bromley) Major shade tree insects, 1940. Jour. Econ.
Ent., 34: (2), 180.

The minute gall insect of white oaks. Scientific Tree Topics 7, Yol.
1, p. 55.

Weather and winter drying. Scientific Tree Topics 7, Yol. 1, p. 50.

236

Journal New York Entomological Society

[VOL. LII

1941. Pruning trees and shrubs. Orange Judd Pub. Co., Inc., N. Y., 237 p.

1942. (With S. W. Bromley) The increasing importance of Coleopterous

borers in shade trees. Jour. Econ. Ent., 35: (2), 169.

1942. Our shade trees. Eevised edition. Orange Judd Pub. Co., Inc., N. Y.,

316 p.

1943. How may the arboricultural profession serve a nation at war? 18th

Natl. Shade Tree Conf., p. 29.

1943. Dutch elm disease control. Scientific Tree Topics 8, Yol. 1, p. 58.
1943. Shelter trees in war and peace. Orange Judd Pub. Co., N. Y., 320 p.

Stanley W. Bromley.

Sept., 1944]

Brown: Butterflies

237

NOTES ON MEXICAN BUTTERFLIES, III, DANAIDiE

By F. Martin Brown

Danainae

124. Anosia berenice Cramer.

G. & S. (1), 1: 3; 2: 638.

Ha. (2), p. 114, pi. 31a.

Ho. (5), p. 662.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 4 ££ 1 2 vi. 16-18. 40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 2 J'J' 2 52 vii.29-viii.-
1.39 (H.H.).

nr. Villagran, Tamanlipas, 1160 ft., 1J iv.28.41 (R.P.).

Hda. Sta. Engracia, Tamanlipas, 1 2 vii.27.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 £ vii.5.36
(H.D.T.).

Jacala, Hidalgo, 4500 ft., 4 J'J' 6 52 vi.25-vii.3.39
(H.H.).

El Sol, Tamazunchale, Sau Luis Potosi, 400 ft., 2 25
v.28-29.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 £ vii.20.39 (H.H.).

nr. Chilpancingo, Guerrero, 1 2 vi.26.41 (R.P.).

El Sabino, nr. Uruapan, Michoacan, 5 J'.J' 1 2 vii.16-
30.36 (H.D.T.).

Chichen Itza, Yucatan, 1 J1 viii.30.36 (H.D.T.).

All this material is probably referable to race strigosa Bates.
A few of the specimens (Villagran, Galeana, Jacala) have the
greyish scales along the nervules of the upper side of the hind
wings so reduced in number as to be difficult to see. There is
some difference in ground color which I am inclined to believe is
related to the freshness of the specimens. Those that are freshly
emerged are brighter fulvous than those that have flown for some
time which are rather dull dilute reddish brown. There is con-
siderable variation in the development of the white spots in the
dark margin of the hind wings. In two cases (a Jacala 2 and an
El Sol 2) these spots are absent, in all of the others the outer row

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Journal New York Entomological Society

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is at least indicated and in some the second row is almost com-
plete. The baso-apical radius of the fore wing varies from 31 to
47 mm. in the males and from 29 to 44 mm. among the females.
There may be a tendency for the more northern specimens to be
brighter and larger, and more clearly strigosa.

125. Anosia cleothera Godart.

G. & S., 1 : 3 ; 2 : 638.

Ha., p. 114, pi. 31b.

Ho., p. 663.

Galeana, Nuevo Leon, 6500 ft., 1 J1 viii.1.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 £ vii.6.36
(H.D.T.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J
v. 29.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 J v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 J' v.10.41 (R.P.).
The Galeana record is rather far north for this species; how-
ever, it has been taken in Texas. In McDunnough’s check list it
is termed eresimus Cramer. I am not sure that cleothera is really
valid for the Central American material, it was described from
‘ ‘ Timor ” ( ! ) and the original description does not apply too well
to our material. Cramer’s species from the Guianas fits it no
better so until a specialist settles the question it is best to let
cleothera stand.

126. Danaus plexippus Linnaeus.

G. & S., 1: 1; 2: 638.

Ha., p. 113 (as archippus F.), Vol. 1, pi. 28c.

Ho., p. 663 (as curassavicce F.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft.. 1 J
v.29.41 (R.P.).

Tancitaro, Michoacan, 6600 ft., 1 2 §5 vii.30-viii.ll.40

(H.H.).

The El Sol specimen and one of the Tancitaro females are f.
fumosa Hulst. Hoffmann lists this species as Diogas curassavicce
Fabricius. Apparently he overlooked Riley’s paper (Tr. Ent.
Soc. Lond., 76: 451. 1929) which definitely linked the Linnean
name with our “Monarch.”

Sept., 1944]

Brown: Butterflies

239

Ithomiinae

My specimens of this subfamily were determined by R. M. Fox,
of the Reading Public Museum. These represent six species of
which Fox says, “All are common members of the Mexican tropi-
cal fauna.’7 All of them were collected by Robert Potts. Of
these six species five are among the twenty-eight Ithomiinae in
Hoffmann’s list. The other has probably been confused by Hoff-
mann with Ithomia patilla Hewitson. I am following Hoffmann’s
order in this listing.

139. Dircenna klugii Hiibner.

G. & S., 1 : 28 ; 2 : 644.

Ha., p. 138, pi. 36e.

Ho., p. 664.

Tuxpango, Yera Cruz, 1500 ft., 1 £ v.8.41.

Fortin, Yera Cruz, 1600 ft., 1 2 v.3-4.41.

Orizaba, Yera Cruz, 2000 ft., 1 2 v.6.41.

141A. Ithomia hippocrenis Bates.

G. & S., 1 : 51, pi. 5, f. 5.

Ha., p. 142.

Ho., p. 665 (I. patilla in part).

Ojo de Agua, Yera Cruz, 1600 ft., 3 3 22 v.12.41.

A pair of these are deposited at the Reading Public Museum.
This species is not listed by Hoffmann.

145. Oleria paula Weymer.

G. & S., 2 : 645 ( L . victoria 1 : 38 in part) .

Ha., p. 150.

Ho., p. 665.

Ojo de Agua, Yera Cruz, 1600 ft., 1 , 1 2 v.12.41.
Hoffmann et al. use the genus Leucothryis for this species.

149. Pteronymia cotytto Guerin.

G. & S., 1: 44; 2: 647.

Ha., p. 155, pi. 40c.

Ho., p. 665.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 3 22
v.28-29.41.

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Tuxpango, Vera Cruz, 1500 ft., 1 2 v.8.41.

Ojo de Agua, Vera Cruz, 1600 ft., 1 J' 2 22 y.12.41.

A female from El Sol and one from Ojo de Agua are deposited
at the Reading Public Museum.

153. Greta oto, Hewitson.

G. & S., 1 : 56 ; 2 : 648.

Ha., p. 164, pi. 41e.

Ho., p. 666.

Ojo de Agua, Vera Cruz, 1600 ft., 2 jy 3 22 v.12.41.
Hoffmann et al. place this and the following species in Hy-
menitis. A pair of oto is deposited at the Reading Public Museum.

154. Greta nero Hewitson.

G. & S., 1: 57; 2: 648.

Ha., p. 163, pi. 41e.

Ho., p. 666.

Fortin, Vera Cruz, 1600 ft., 1 y v.3.41.

SATYRIDiE

As is so frequent with collections made by inexperienced col-
lectors in the tropics the catch of Satyridse brought back by both
Hoogstraal and Potts is small in comparison with what it might
have been. The bulk of the specimens taken are Euptychia.
Most of the jungle-loving species were not captured.

159. Tisiphone maculata Hopffer.

G. & S., 1 : 72, 651.

W. (3), p. 184, pi. 44b.

Ho., p. 666.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 jy
1 2 v.28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 22 vi.26-
28.40 (H.H.).

The El Sol female is fresh ; all of the other specimens are worn.

163. Taygetis virgilia Cramer.

G. & S., 1 : 97, 659.

W., p. 187, pi. 45a.

Sept., 1944]

Brown: Butterflies

241

Ho., p. 666.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 jy
v.28-29.41 (R.P.).

167. Taygetis inconspicua Draudt.

Ho., p. 667.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 7 jy
iv.30, vi. 28-29. 41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 y vi. 28.40

(H.H.).

El Pujal, San Luis Potosi, 100 ft., 2 yy vi. 17-18. 39
(H.II.).

The El Banito male and several of the May specimens from
El Sol are worn ; the others are reasonably fresh. I suspect that
this is nothing more than a form of virgilia. The genus will be
treated in monographic form as a member of my series of papers
devoted to Ecuadorian butterflies.

172. Neonympha gemma Hiibner.

G. & S., 1 : 92, 657, pi. 8, f. 12.

W., p. 223 (as Cornelius F.), pi. 49f.

Ho, p. 667.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft, 2 jy 1 ? vi. 18-20.40 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 2 yy vii.6.36
(H.D.T.).

Jacala, Hidalgo, 4500 ft, 3 jy vi.24-29.39 (H.H.).
Fortin, Vera Cruz, 1600 ft, 1 y v.4.41 (R.P.).

179. Megisto hesione Sulzer.

G. & S, 1 : 75, 651 (as ocirrhoe).

W, p. 194, pi. 46d.

Ho, p. 667.

El Sol, Tamazunchale, San Luis Potosi, 400 ft, 7 yy
iv. 30-31, v.28-29.41 (R.P.).

Ojo cle Agua, Vera Cruz, 1600 ft, 2 55 v.12.41 (R.P.).

181. Megisto mollina Ilewitson.

G. & S, 1: 76, 652, pi. 8, ff. 13, 14 (fig. as westwoodi).

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[Vol. LII

W., p. 195, pi. 46e.

Ho., p. 668.

El Sol, Tamazunchale, San Lnis Potosi, 400 ft., 5 J'J'
v.28-29.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 3 J'.J' 3 55 v.12.41
(R.P.).

The females and one male from Ojo de Agua are f. westwoodi
Butler.

182. Megisto fetna Butler.

G. & S., 1 : 77, 652, pi. 8, ff. 15, 16.

W., p. 195.

Ho., p. 668.

Chilpancingo, Guerrero, 1 vi. 26.41 (R.P.).

184. Megisto terrestris Butler.

G. & S., 1 : 79.

W., p. 200.

Ho., p. 668.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 1 S vi.19.40 (H.H.).

This specimen is probably not true terrestris which hails from
eastern South America but may be the same form regarded as
terrestris by Hoffmann. It is far north of Hoffmann’s record
“Selva virgen de Campeche (Tierra caliente).” It compares
well with Weymer’s figure noted above.

188. Megisto rubricata Edwards.

G. & S., 1 : 82, 655, pi. 8, f . 5.

W, p. 202.

Ho., p. 668.

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1000 ft.,
1<? vi.14.40 (H.H.).

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1600
ft., 1 vi.19.40 (H.H.).

Jacala, Hidalgo, 4500-5100 ft., 3 J'J' 2 5? vi.24.39

(H.H.). ^

Sept., 1944]

Brown: Butterflies

243

190. Megisto renata disaffecta Butler.

G. & S., 1 : 82, 655, pi. 8, f. 5 (as renata).

W, p. 204, pi. 47d.

Ho., p. 668.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 y 1 2
iv.30-31.41 (R.P.) .

Fortin, Vera Cruz, 1600 ft., 1 y v.3.41 (R.P.).

Tuxpango, Vera Cruz, 1500 ft., 1 y v.9.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 3 yy v. 12.41 (R.P.).

El Sabino, nr. Uruapan, Michoacan, 1 ^ 1 J vii.15-
30.36 (H.D.T.).

Chichen Itza, Yucatan, 2 yy viii. 30.36 (H.D.T.).

Determination tentative.

191. Megisto hermes Fabricius.

G. & S., 1 : 86, 656, pi. 8, ff. 6, 7 (as camerta).

W., p. 207, pi. 48a.

Ho., p. 668.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon,
1600 ft., 3 yy vi. 17-18.40 (H.H.).

Monterrey, Nuevo Leon, 1800 ft., 2 yy 1 2 iv.27.41
(R.P.).

nr. Villagran, Tamaulipas, 1160 ft., 1 y iv.28.41 (R.P.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 yy
2 52 v.28-29.41 (R.P.) .

El Banito, Valles, San Luis Potosi, 200 ft., 3 yy
vi. 27.41 (H.H.) ; 1 y iv.30.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 2 yy 5 22 vii.17-
21.39 (H.H.).

Hda. Potrero Viejo, Vera Cruz, 1500 ft., 1 2 v.5.41
(R.P.).

Tuxpango, Vera Cruz, 1500 ft., 1 2 v.9.41 (R.P.).

Fortin, Vera Cruz, 1600 ft., 2 52 v.4.41 (R.P.).

Ojo de Agua, Vera Cruz, 2000 ft., 2 yy 1 2 v. 12.41
(R.P.).

Orizaba, Vera Cruz, 2000 ft., 4 yy 2 22 v.6.41 (R.P.).

60 mi. So. of Victoria, Tamaulipas, 1 y 1 2 vii.6.36
(H.D.T.).

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Journal New York Entomological Society

[Yol. lii

El Sabino, nr. Uruapan, Michoacan, 5 4 52 yii.15-

30.36 (H.D.T.).

Chichen Itza, Yucatan, 2 55 viii.30.36 (H.D.T.).

An extremely varied series. The species “kermes” may be
composite.

197. Megisto libye Linnaeus.

G. & S., 1 : 83, 655.

W., p. 212, pi. 48e.

Ho., p. 669.

Ojo de Agua, Vera Cruz, 1600 ft., 1 5 v.12.41 (R.P.).

198. Megisto glaucina Bates.

G. & 1 : 90, pi. 8, if. 18, 19.

W, p. 216, pi. 49b.

Ho., p. 669.

Ojo de Agua, Vera Cruz, 1600 ft., 1 (J 1 5 v.12.41
(R.P.).

Megisto sp.

Fortin, Vera Cruz, 1600 ft., 1 v.4.41 (R.P.).

Ojo de Agua, Vera Cruz, 2000 ft., 1 v.6.41 (R.P.).

A species near nebulosa Butler. Differing from that South
American species in having five ocelli on the under side of the
fore wing.

200. Pindis squamistriga Felder.

G. & S., 1 : 80, 654, pi. 8, f. 24 (fig. as zabdi).

W., p. 224, pi. 50a.

Ho., p. 669.

Jacala, Hidalgo, 4500 ft., 2 vi.23-29.39 (H.H.).

El Sabino, nr. Uruapan, Michoacan, 1 (5^^.15-30.36

(H.D.T.).

Sta. Lucretia, 2 <5<5 ix.4.39 (H.D.T.).

The two Jacala specimens are badly battered; fortunately one
entire fore wing is intact on one specimen, sufficient to recognize
the genus and species. This is the first record of the species from
so far northeast in Mexico.

Sept., 1944]

Brown: Butterflies

245

202. Paramecera xicaque Reakirt.

G. & S., 1 : 101, 660.

W., p. 225, pi. 50a.

Ho., p. 669.

Tancitaro, Michoacan, 6600 ft., 1 J1 vii.25.40 (H.H.).

A battered specimen.

BRASSOLIDiE

216. Opsiphanes cassina fabricii Boisduval.

G. & S., 1 : 127, 664 (as cassice).

F. (4), p. 302, pi. 51d.

Ho., p. 671.

El Banito, Valles, San Luis Potosi, 200 ft., 3 1 5

vi.28.40 (H.H.).

Hoffmann reports this species from both coasts as far north as
Sinaloa and Tamanlipas in contrast to Fruhstorfer’s report of it
only as far north as Tepic, Nayarit on the west coast and no men-
tion of it in eastern Mexico.

218. Eryphanis aesacus aesacus Herrick-Schaffer.

G. & S„ 1 : 137, 666.

F. , p. 313, pi. 64b.

Ho., p. 671.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 J'J'
1 2 v.28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 $ v.28.40
(H.H.).

El Pujal, San Luis Potosi, 100 ft., 2 vi.18.39

(H.H.).

These records extend the range of the species a little further
north than that given by Hoffmann. Fruhstorfer says of the
species, 1 1 Everywhere rare. ’ ’

MORPHID.ZE

225. Morpho peleides montezuma Guenee

G. & S., 1 : 119, 663 (as peleides).

F., p. 344.

Ho., p. 672.

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Journal New York Entomological Society

[Vol. LII

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 5 J'J'
y.29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 J'J'
vii.22.39, vi. 26.40 (H.H.).

Ojo de Agua, Vera Cruz, 1600 ft., 3 J'J' 1 § v.12.41
(R.P.).

The Ojo de Agua material is in very poor condition, especially
the female. The other specimens are in good condition. The
records for San Luis Potosi extend the range of the species a
little to the north.

BIBLIOGRAPHY

1. Godman & Salvin, Biologia Centrali- Americana, Rhopalocera, 1 : 1-139.

1879-1881.

2. Haensch, in Seitz’ Macrolepidoptera of the World, vol. 5, p. 113-171.

1909- 1910.

3. Weymer, in Seitz’ Macrolepidoptera of the World, vol. 5, p. 173-283.

1910- 1912.

4. Fruhstorfer, in Seitz’ Macrolepidoptera of the World, vol. 5, p. 285-356.

1912-1913.

5. Hoffmann, Anales del Instituto de Biologia (Mexico), 11: 639-739.

1940.

Sept., 1944]

Ginsburg: Mosquitoes

247

OUTDOOR PROTECTION FROM MOSQUITOES*

Joseph M. Ginsburg

Biochemist in Entomology, New Jersey Agricultural
Experiment Station

Even in areas where mosquito control work has been systemati-
cally conducted for many years it has not been possible, with our
present means and methods, completely to eradicate the mosquito.
Under conditions favorable for its rapid development, such as
during summers of abundant rainfall, it becomes at times a seri-
ous interference with human comfort.

When a female mosquito bites or pierces the skin to suck our
blood she injects into the wound a small amount of a poisonous
substance, the chemical composition of which has not been defi-
nitely established. The effect of this injected material varies
with different species of mosquitoes as well as with the suscepti-
bility of the bitten individual. While a small number of people
appear to possess a certain degree of immunity, either natural or
acquired, against this poison, the great majority of us, especially
children, suffer from its effects. Immediately following the bite,
an itching sensation is felt which may be followed by considerable
swelling. The scratching induced by the irritation may cause a
secondary blood infection, especially among children, leading to
serious results. The irritation may be relieved by washing the
swelling with any one of the following solutions : soft soap, alco-
hol, glycerin, iodine or borax.

Complete elimination of mosquitoes indoors can be readily
accomplished by proper screening, spraying, or fumigating. On
the other hand, protection from mosquito annoyance outdoors
constitutes a difficult problem, the solution of which cannot
always be successfully attained.

Mosquito repellents. — -Various chemicals, possessing repelling
properties against mosquitoes, have been recommended and com-
mercialized. They are applied in form of lotions, ointments or
powders to the exposed parts of the body. The duration of the

* Journal Series paper of the New Jersey Agricultural Experiment Station,
Rutgers University, Department of Entomology.

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Journal New York Entomological Society

[Vol. LII

protection may vary from a short time to several hours, depend-
ing on the inherent repellent properties of the chemical, thorough-
ness of application, species and density of the mosquito popula-
tion, degree of attractiveness possessed by the individual, and
atmospheric factors.

Citronella is one of the oldest mosquito repellents used either
in its natural liquid state or in combination with various other
compounds. One of the early formulas recommended by Dr.
L. 0. Howard (1923) contains the following ingredients: oil of
citronella 2 ounces, spirits of camphor 2 ounces, and oil of cedar
1 ounce. This preparation can also be made in form of a salve,
if desired, by mixing in enough petrolatum or similar semisolids.
One thorough application may remain effective for 1 or 2 hours.
Dr. Herms (1939) finds the following formula successful: citron-
ella 3 parts, kerosene 2 parts, and coconut oil 4 parts. To this
mixture is added 1 per cent carbolic acid. Within the last few
years longer lasting and more efficient repellents have been syn-
thesized, as a result of research conducted at the Federal Bureau
of Entomology in Washington and at Rutgers University (Gran-
ett 1940), in cooperation with various commercial concerns.
Three of these are at present widely used, namely, dimethyl
phthalate, Indalone, and formula No. 612.

Though mosquito repellents are contributing a great deal
toward relief from mosquito annoyance to individuals, such as
night watchmen, military pickets, mosquito workers and fisher-
men, compelled to remain exposed for considerable lengths of
time in mosquito-infested areas, their frequent application en-
counters many drawbacks. First, not all the repellent mixtures
thus far developed repel mosquitoes from a distance. In virtu-
ally every case the mosquito has to alight, or at least come very
close to the treated surface, before the chemical acts on it. Com-
plete and thorough coverage is, therefore, necessary, otherwise
untreated spots may be bitten. Second, virtually all of the repel-
lents cause sharp smarting on delicate parts of the skin such as
eyelids, face and forehead. This irritation is apt to become
rather pungent and often unbearable to many sensitive indi-
viduals under warm, humid atmospheres and other conditions
when copious perspiration occurs. Third, because of the solvent

Sept., 1944]

Ginsburg: Mosquitoes

249

properties of the repellents, a treated individual must avoid
coming in contact with varnished and painted surfaces. Either
paint or varnish will be partially removed, resulting in staining
of clothes or any other objects incidentally touched. Fourth, the
repellent does not protect a group, much less a large outdoor
audience, unless each individual is treated. Fifth, continuous
feel of a greasy, somewhat irritating solvent all over one’s skin
becomes rather unpleasant. Unless compelled by military duty
or night work, few individuals will willingly submit themselves
to this treatment night after night. Instead they would rather
stay indoors or in a well-screened porch.

Mosquito sprays. — In view of these objectionable features, the
writer became interested in developing a mosquito spray which
should, with no injury to man, animals, and plants, completely
free an area from adult mosquitoes where groups of people could
spend a summer evening with no mosquito annoyance and with
perfect body comfort. This effort coincided with another prob-
lem, that of developing a mosquito larvicide which should kill
mosquito larvae and pupae with no injury to fish, water fowl, and
aquatic plants, offering at the same time no fire hazard. Such
a product was called for in residential sections, ornamental ponds,
game conservation parks, and similar places where mosquito oil
was objectionable. By 1931 the New Jersey Py rethrum Mosquito
Larvicide was developed and came into wide use as a substitute
for oil wherever the latter meets with objections, Ginsburg
(1930). The larvicide is essentially an emulsion consisting of
66 per cent kerosene, 0.5 per cent sodium lauryl sulfate as emulsi-
fier, 0.07 per cent pyrethrins, and about 34 per cent water. The
concentrated stock emulsion is mixed just before spraying with
about 10 parts of clear water, which may be taken directly from
• the area to be sprayed.

Preliminary tests conducted during 1934 (Ginsburg 1935) indi-
cated that spraying with this diluted larvicide might also prove
effective in ridding a given area from adult mosquitoes. Further
extensive experiments carried out during 1935-36 (Ginsburg
1936, 1937) by the writer in cooperation with the various County
Mosquito Commissions in New Jersey have substantiated this
assumption and have established a definite method of procedure
for spraying.

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Journal New York Entomological Society

[Yol. lii

The experiments during these three years have given us some
fundamental knowledge as well as authentic information concern-
ing the possibility and practicability of eliminating mosquito
annoyance from outdoor public gatherings without in any way
interfering with human comfort and activities. We have learned
how to accomplish this relief without appreciable injury to plant
life. These results have also thrown some light on the physical
and chemical mechanism involved as well as the role that each
ingredient of the larvicide plays in this process. From the experi-
ence thus far gained the following principles may be evolved as
guides in this work.

In order successfully to eliminate adult mosquitoes from a
given area outdoors, two objectives must be attained. First, all
female mosquitoes resting in the grass, shrubs, and throughout
the area to be treated must be killed. This is accomplished by
thoroughly spraying the entire grounds before the audience
gathers. Second, the influx of mosquitoes from the surroundings
into the protected area must be prevented. For this purpose the
spray is directed upward, as high as the pressure permits, so as
to saturate the atmosphere with a fine mist of larvicide. This
air-fogging may have to be repeated two or three times during the
affair, depending on the species and density of mosquitoes, the
flight intensity, the direction and velocity of wind, and other
atmospheric factors. It should be emphasized that, for complete
protection, it is just as important to kill or incapacitate all the
female mosquitoes hiding in vegetation as it is to bar those in the
surroundings from flying in. The spray should be applied in
highly dispersed form, as a fog or mist. The finer the liquid
particles, the less will be the danger of injury to plants and the
longer will it remain floating in the air. The diluted larvicide
must be kept thoroughly mixed during spraying.

The mechanism involved in this process is twofold, repellency
and contact killing. The female mosquito when in contact with
the larvicide is either killed or paralyzed to such a degree that,
for an indefinite period, she can neither fly nor bite. These symp-
toms are brought about primarily by the pyrethrins incorporated
in the larvicide. The petroleum oil acts as a carrier for the
pyrethrum. It should be mentioned here that neither an aqueous

Sept., 1944]

Ginsburg: Mosquitoes

251

spray of pyrethrum extract nor a kerosene emulsion when applied
separately gave satisfactory protection. Evidently the combi-
nation of both ingredients is necessary for adequate protection.
The water in the larvicide merely serves as the outer phase in
which the two toxic ingredients are dispersed by the aid of the
emulsifier.

The question has been asked : What happens to the mosquitoes
in the sprayed area? The following observations may offer an
explanation. When tents were sprayed inside with the larvicide,
some mosquitoes were found on the ground apparently dead while
others remained on the canvas in a quiescent condition. When
disturbed they attempted to fly but soon again attached them-
selves to the canvas. They remained for some time in this partly
paralyzed condition and finally either dropped to the ground or
flew away. These symptoms closely resemble those observed in
other insects treated with pyrethrum insecticides and can, there-
fore, be attributed directly to the pyrethrins incorporated in the
larvicide. Again, while checking the mosquito density the writer
observed on several occasions that when mosquitoes reach the
treated area they abruptly change their course of flight, turning
back or at a right angle. This behavior suggests repellency.
Evidently mosquitoes find the oil, the pyrethrum, or both com-
bined so disagreeable that they keep away from the treated area.

DIRECTIONS FOR SPRAYING

The concentrated larvicide is first well shaken or stirred. It is
then mixed with 12 parts of water (1 quart to 3 gallons) in the
sprayer. This diluted mixture is ready for spraying. During
the spraying operation it should be frequently shaken or stirred
in order to insure uniform distribution of the larvicide.

Procedure. — Before the affair starts, the entire area, including
grass, shrubs, bleachers, sheds, benches, or any other place where
mosquitoes may rest during the day, is thoroughly sprayed with
the diluted larvicide. This should kill all mosquitoes as well as
many other insects hiding in the grass, shrubs, and other places.
The next object is to prevent mosquitoes from the surroundings
from coming into the protected area. For this purpose the spray
is directed upward so as to saturate the atmosphere with a fine
mist or fog of the larvicide. If the spray is applied against the

252

Journal New York Entomological Society

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wind the fine mist or fog will drift with the wind throughout the
area. This entire operation should be finished before the gather-
ing takes place. Just about dusk or when the mosquitoes from
the outside begin to fly in, another fogging is necessary. For this
purpose the spray is applied as high as the sprayer permits and
primarily on the side from which the wind is blowing. This mist
in the air wards off mosquitoes from outside the treated area.
If no noticeable wind prevails it may be necessary to fog all
around the area, directing the spray upward so as to keep the
inflying mosquitoes away. A thorough fogging about the time
when the mosquitoes start coming in should be sufficient for the
rest of the evening. Under very heavy infestations, where the
mosquitoes are coming in large numbers, a second fogging about
9 : 30 may be necessary.

Apparatus Required. — (1) on small areas, such as backyards,
porches, and private lawns, a garden sprayer, knapsack sprayer,
bucket pump, or electric sprayer capable of producing a fine
spray, about 10 or 15 feet high, may be used with success.

( 2 ) For spraying large areas, such as picnic grounds, stadiums,
and open air theaters, a power sprayer capable of developing a
pressure of 200 pounds or more per square inch and equipped
with a spray gun is necessary. The larvicide has been found
most efficient in protecting outdoor audiences on comparatively
large areas where power sprayers can be used.

Precautions. — Before attempting to treat an area, one should
make a thorough survey in order to ascertain the following
points: sources of mosquitoes; possible mosquito resting places;
direction of wind; type of vegetation present; water supply for
mixing the larvicide ; kind of spraying outfit necessary ; nature of
gatherings; and any difficulties that may be anticipated during
the spraying operation. This information should enable the
operator to plan his method of procedure.

PRESENT STATUS OF THE NEW JERSEY PYRETHRUM
MOSQUITO LARVICIDE

Since this spray was introduced, numerous outdoor evening
concerts, carnivals, church parties, community gatherings, and
lawn parties have been fully protected from mosquitoes either
directly by the County Mosquito Commissions at a nominal cost,

Sept., 1944]

Ginsburg: Mosquitoes

253

or indirectly through their assistance, guidance, and advice. The
number of persons present on these occasions varied from small
groups to many thousands. Records show that in several in-
stances some 20,000# persons in one gathering have enjoyed an
open air evening concert with no mosquito discomfort as a result
of spraying the stadium where the affair was held. When the
larvicide was sprayed as directed, no injury has resulted to grass,
shrubs, trees, ornamental plants or aquatic plants. Up to and
including 1942, its use increased with each successive season.

Prior to 1934 most of the pyrethrum used in this country came
from Japan, and small amounts from Dalmatia and the Kenya
Colony in British East Africa. By 1939 the picture was re-
versed; most of our pyrethrum was imported from Kenya and
very little from Japan. When World War II broke out, cultiva-
tion of pyrethrum in the British possessions, for various reasons,
decreased. Shipping shortage and submarine losses still further
curtailed the flow of pyrethrum flowers to this country. After
Pearl Harbor, practically all the available and potential supplies
of pyrethrum were taken over by the United States Government
for army use. At present the War Production Board does not
allow the use of pyrethrum for preparing the larvicide. Thus,
the excellent record of this newly developed method for tempo-
rary relief from biting female mosquitoes has been suddenly
terminated for the duration or until enough pyrethrum becomes
available to supply a surplus, above the amount necessary for our
armed forces.

Intensive testing is now being conducted with various other
chemicals as possible substitutes for pyrethrum. At this date,
however, none has proved equal to pyrethrum from the stand-
points of effectiveness against the mosquito and of safety to man,
animals, and plants.

LITERATURE CITED

Ginsburg, J. M. 1930. Studies of pyrethrum as a mosquito larvicide. Proc.
17th Ann. Meet. N. J. Mosq. Ext. Assoc., p. 57.

. 1935. Larvicides and method for temporary protection from adult

mosquitoes in limited areas. Proc. 22nd Ann. Meet. N. J. Mosq.
Ext. Assoc., p. 147.

* Essex County Symphony Society, at the Newark School Stadium.

254

Journal New York Entomological Society

[Yol. lii

. 1936. Protection of outdoor gatherings from the mosquito pest.

Proc. 23rd Ann. Meet. N. J. Mosq. Ext. Assoc., p. 166.

. 1937. Principles underlying the protection of outdoor meetings

from the mosquito pest and methods of application. Proc. 24th
Ann. Meet. N. J. Mosq. Ext. Assoc., p. 5.

Granett, P. 1940. Studies of mosquito repellents. Jour. Econ. Ent.,
33(3) : 563-71.

Herms, W. B. 1939. Medical Entomology. MacMillan, New York.
Howard, L. O. 1923. Remedies and preventives against mosquitoes.
U. S. D. A. Farmers’ Bull. No. 444.

Vannote, R. L. 1937. Methods of applying the larvicide as a repellant.
Proc. 24th Ann. Meet. N. J. Mosq. Ext. Assoc., p. 11.

Sept., 1944]

Clench: Lyoenhle

255

NEW NEOTROPICAL THECLINZE (LEPIDOPTERA,
LYC^NID^E)

By Harry K. Clench
Cambridge, Mass.

The following new Theclinge have come to my attention recently,
two of them during the arranging of the Neotropical Lycsenidse
in the collection of the Museum of Comparative Zoology (M.C.Z.)
and one in a loan of study material from the American Museum
of Natural History.

Thecla caramba, new species

Eyes ringed with pale, green. Frons metallic green. Collar above obscure,
laterally brown. Palpi rusty gray, terminal sediment dark. Antennce
black above, white annulate below. Thorax above black-brown (scaled with
metallic blue?), covered anteriorly, laterally and posteriorly with long, back-
directed hairs, bluish in color; below with moderately long, red-brown hair.
Abdomen above metallic blue, becoming gray towards the anal extremity;
below pale yellow, or rich cream. Legs largely black, ringed narrowly and
infrequently by white.

Upperside :

Male. Both wings dark, lustrous blue, with a purple luster. Fore wing
with a narrow outer marginal border, thickening slightly at the apex. Find
wing with costa gray-brown, and inner margin gray. Outer margin very
narrowly black. Anal lobe rusty. Outer margin very slightly produced at the
vein-ends from Mx to the anal angle, but no indication of any tails is present.
Fringe of both wings brown, on hind Aving paler outwardly between the veins.

Underside:

Male. Both wings bright pea-green. Fore wing Avith a gray area from Cu2
to the inner margin. The green along this gray area is bluish, basad. On
the outer margin the green extends down to 2A. Find wing with a white
spot on the costa, near outer angle, basally edged with dark red. A similar,
but post-discal, spot in M2-M3 and one in Cuj-Cua. In Cu2-2A is a white
dash, basally edged with red, and in 2A-inner margin is a similar, but heavier
dash. Anal lobe red, extending on outer margin to Cu2. In Cm-Cua is a
submarginal red dash. Fringe of both wings rusty-red, outwardly paler on
the hind Aving betAveen the veins.

Length of fore wing : Male, 15 mm.

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Journal New York Entomological Society

[Vol. LII

Holotype, male, Massaranduba-Blumenau, Brasil (ex coll. E. I.
Huntington (no. 1009)); in the American Museum of Natural
History.

Remarks. Evidently closely allied to amyntor Cramer,1 of
which, unfortunately, no males have been examined. The holo-
type of caramba was compared with a female (in M.C.Z., no data,
ex coll. A. G. Weeks) of amyntor , the description and figure of
Draudt (1919, in Seitz, Macrolep. World, 5, p. 762, pi. 153 i),
and Cramer’s descriptions and figures of amyntor ( loc . cit.) and
menalcas (1782, Pap. Exot., 3, p. 117, pi. 259, figs. A and B).

Above, caramba differs from amyntor (of Draudt) in the nar-
rower, better defined marginal border. Below, from the female
at hand, the present species seems to have more of an anal lobe
and less prominent white markings. The absence of a tail is
probably very significant, and would seem to connect amyntor
to the acaste group, particularly to those members of that group
with a green frons. It certainly destroys Draudt ’s character for
splitting the “ amyntor -group” into two sections ( i.e ., the tailed
section, comprising the true amyntor group, to which caramba
obviously belongs, and the tailless section, more aptly called the
acaste group, and to which caramba bears little (relative)
resemblance.

Thecla punona, new species

Eyes ringed with white. Frons with moderately profuse, long, erect pale
gray hairs. Collar above with brown and gray hairs intermingled; on the
sides, white. Antennce largely white below, black above, but with thin an-
nulations of each extending into the other; club black. Thorax black above,
with sparsely strewn bronzy-brown hairs; below with pallid gray hair, quite
profuse. Abdomen gray-tan above, gray below.

Upperside :

Female?. Both wings slightly lustrous brown. Hind wing with outer
margin from M3 to the anal angle scalloped, and with a narrow whitish
marginal thread-line from Cu2 to 2A. Fringe of both wings dirty white,
basally darker ; that of hind wing obscurely dark at the vein-ends.

Underside :

Female?. Fore wing tan. A post-discal, slightly curved, dark brown line,
outwardly white, proceeds from costa to Cu2. Immediately without this line
at the apex is a hoary area, extending from costa roughly to M3, A sub-

1 1779, Pap. Exot., etc., 1, p. 76, pi. 48, fig. E.

Sept., 1944]

Clench: Lyc^enid^e

257

marginal, frequently indefinite line crosses faintly through this, increasing in
visibility below it until its disappearance at 2A. Kind wing hoary gray,
darker on the outer margin. A submarginal row of faint yellowish or pallid-
fulvous lunules parallels entirely the outer margin. A curved, irregular, dis-
cal, brown line, tan within and white without, crosses from costa to inner
margin, and a post-basal, similar one also, though necessarily shorter. The
two close off an area, slightly darker than the surrounding ground, which is
just basad of the Center. Occasionally the ground color is so disposed as to
give the appearance of a row of pale gray lunules capping the yellowish ones
on the outer margin. Fringe similar to the upperside but rather darker.

Length of fore wing : Female?, 9.5-11 mm.

Holotype, female (f), Puno, Peru, 12,500 feet, Nov. 1, 1898
(Wm. J. Gerhard).

Paratypes, two females ( ?), same data as holotype.

Holotype and one paratype, no. 26258 in the M.C.Z. One para-
type in the collection of the author.

Remarks. Rather closely allied to oribata Weymer2 and pos-
sibly only subspecifically distinct from it. The principal differ-
encse are : the less apically produced fore wing and less anally
produced hind wing (if Weymer ’s figure is correct); the less
distinct outer row of spots on the fore wing below, and the less
apparent submarginal line on the hind wing below; the more
prominent basal dashes ; the less distinctly black-checked fringe,
particularly on the hind wing. A single specimen from Chulu-
mani, Bolivia (Nov. 28, 1898, W. J. Gerhard), not made part of
the type series, approaches oribata in several of these respects.

Weymer ’s figure shows the lines on the underside to be quite
dark, almost black, while in punona they are brown. It is prob-
able, however, that the figure errs somewhat in this respect.

Draudt (1919, in Seitz, Macrolep. World, 5, p. 760, pi. 153 h)
described a new species under the name of anosma, which he later
(Seitz, add. and corr., p. 823) synonymized to oribata. In his
description he mentioned “incoherent dark lunular spots” and
“longer small tail on the lower median vein” neither of which is
particularly clear. The latter particularly would seem to exclude
it entirely from the culminicola-gr owp. It is presumed that by
“median ’ ’ he meant 1 1 cubital. ’ ’

2 1890, Lepidopteren gesammelt . . . von A. Stiibel (Rhopaloeera by Wey-
mer), p. 123, pi. 4, fig. 8.

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Journal New York Entomological Society

[Yol. LII

Thecla kalikimaka, new species

Thecla hemon: Hoffman, 1940, Anales Inst. Biol. Mex., 11, p. 704 (no. 586)
( nec hemon Hiibner). _ Ciu. i w >

Eye ringed obscurely with white. Frons brown. Collar brown. Antennce
black-brown, club dull fulvous ; below, white annulate, club brighter fulvous.
Palpi brown, slightly hoary within. Thorax metallic green-blue above,
slightly hairy; below, covered with uniform gray-brown hair. Abdomen above
green-blue, below brown. Legs brown-black, tarsal segments white annulate.

Upperside :

Male. Both wings somewhat bronzy green. Fore wing with a compound
scent-pad consisting of a dark dash across the cell-end and a rectangular
brown patch just beyond. Outer margin moderately heavily edged with
brown, thickening considerably towards the apex. The basal limit of this
brown is very tenuous and indefinite. Hind wing with a small anal area of
brown, and two tails, one each at Cux and Cu2, the latter longer and emitted
tangentially. Anal lobe deeply incised, as is usual in this group.

Underside :

Both wings brown. Fore wing with a faint, dull, metallic green iridescence.
Paler on inner margin. A discal dark brown line crosses from costa to Cu2,
quite straight. Mind wing with a straight discal line from Rs opposite the
outer angle to just beyond Cu2 in the center of the wing. From there it be-
comes metallic green, and proceeds to 2A, where it angles sharply basad and
terminates at the inner margin. A submarginal line starts at about M2 and
proceeds parallel to the outer margin, reaching just beyond Cu2 where it angles
basad, to terminate at the inner margin, edged inwardly from Cu2 with a few
metallic scales. A pale, hardly noticeable marginal line runs from outer
angle to apex, becoming very obscurely greenish from Cu± on. Anal region
restrictedly irrorated with pale scales. A faint line, post-basal, starts on
costa and proceeds to lower cell-end. Fringe of both wings brown.

Length of fore wing : Male, 16 mm.

Holotype, male, Jalapa, Mexico, date and collector unknown,;
ex coll. A. G. Weeks, Jr.

Paratype, male, same locality, Sept. 8, 1884 (“Morrison”),
ex coll. F. A. Eddy.

Holotype, M.C.Z. 26257. Paratype in the author’s collection.

Remarks. Apparently most closely allied to mavors and
triquetra (references and authorships of these names below).
From both it differs in the brown color of the underside, which
in these species is bright metallic green or blue. The lines below
are less apparent than in either of the two species (except thaf
in kalikimaka there is a discal line on the fore wing, lacking in

Sept., 1944]

Clench: Lyc-enid^e

259

mavors). Above it differs from triquetra in the green color (blue
in that species), and from mavors in the lack of a pale, almost
white, anal line. The apical and marginal border of the fore
wing appears slightly heavier than in either mavors or triquetra.
It would seem closest to mavors, since it has the two tails of that
species, as opposed to the single one of triquetra.

A brief discussion of the species of this group (as limited by
Drauclt, p. 750) might not be amiss.

Thecla mavors Hiibner (1818, Zutr. zur Samml. Exot. Schmett.,
1st. Hund., p. 31 (no. 95), figs. 189, 190). Quite variable, but
whether racially or not cannot be determined from the material
at hand. The male is metallic green above, with a compound
scent-pad, and is bordered narrowly and indistinctly with black-
brown on the fore wing. Hind wing at the anal angle edged
with very pale green, almost white. Below green, fore wing
brown on inner margin, otherwise practically immaculate. Hind
wing with a discal and submarginal line, each angled basad at
Cu2-2A. Outward of the latter is a band of brownish maroon
from Mi to the anal angle. Anal lobe similarly colored. Two
tails, the larger at Cu2 tangentially emitted.

Female, brown above and below. The latter surface crossed
on both wings by thin brown lines; two on fore wings, parallel
to the outer margin, and a short one crossing the cell-end. An-
other pair on hind wing, also parallel to the outer margin, but
angling basad at Cu2-2A. A post-basal line commences on costa,
crosses cell-end and stops.

Specimens in the M.C.Z. from Suapure, Venezuela; Cusilluni,
Bolivia; Cumato Arepo, Savanna, Trinidad; Rio Tapajoz, Brasil.

A Suapure female has the two lines on the hind wing abnor-
mally close together, but whether or not this is an individual
variant cannot be stated without additional material.

Thecla triquetra Hewitson (1862-1869, 111. Diurn. Lep. Lycas-
nidas, p. 76, pi. 28, figs. 17, 18, 19). Differs from mavors in the
male by being blue above, having but one tail (also tangentially
emitted), the bluer color below, and the presence of a discal line
on the fore wing, and heavier ones on the hind wing. Females
differ from mavors females chiefly in the lack of a Cui tail.

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Journal New York Entomological Society

[Vol. LII

Otherwise they appear very similar. There is, perhaps, a slight
difference in the shape of the hind wing, but more specimens are
needed to confirm it.

In the M.C.Z. from Blumenau and Rio de Janeiro, Brasil.

Theda pawper a Felder (1865, Reise Novara, Lep. 2, p. 246,
pi. 31, fig. 15). May be distinguished from the above in the
male by the dark green color and the lack of a scent-pad on the
fore wing. Differs below in that the inner of the two longest
lines touches the outer at the anal angle.

Apparently not in the M.C.Z. (see under ella).

Theda drucei Lathy (1926, Ann. Mag. Nat. Hist., (9) 17, p.
41). This species, also not in the M.C.Z., is figured in the Trans.
Ent. Soc. London, 78, pi. 9, fig. 9 (plate accompanying paper
that begins on p. 133). It apparently belongs near pauper a and
harrietta, but differs, in the male, by the possession of a scent-pad.
There are other differences as well.

Theda ella Draudt (1919, in Seitz, Macrolep. World, 5, p. 750,
pi. 148 b as mavors). Differs from pauper a in that the inner
line below on the hind wing is w-shaped at the anal angle. This
is. the difference brought out by Draudt. He noted the fact that
paupera was rare, and very likely based this difference on the
figure of Felder, which does show no “W” at the anal angle
of the hind wing below. This character, however, may be very
faint (as in the single specimen here provisionally assigned to
ella ), and consequently overlooked when Felder’s specimen was
figured.

A single specimen in the M.C.Z. (Bogota, Colombia?) ex coll.
A. G. Weeks, may possibly be this species. It has, however, much
heavier borders above than Draudt ’s illustration shows, and
narrower lines below. It might also be paupera , but again the
markings below are all thinner than Draudt ’s and Felder’s
illustrations show.

Theda harrietta Weeks (1901, Can. Ent., 33, p. 294). This
is a perfectly good species, allied to paupera and ella. Besides
the greatly differing color above, the lines below are rather dif-

Sept., 1944]

Clench: Lyozenhle

261

ferent, the central apex of the “W” being rather shorter than
in the specimen cited above, and in Draudt’s figure of ella. The
discal line on the fore wing extends into the Cu2-2A interspace,
which it apparently does not in the other species. In other
Theclinae, however, this character has been found to be individu-
ally variable.

One specimen in the M.C.Z. (Type, M.C.Z. no. 16673) from
Coroico, Bolivia, April 20, 1899 (Wm. J. Gerhard).

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Journal New York Entomological Society

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NOTE ON THE DEATH-FEINT OF BRUCHUS
OBTECTUS (SAY)

The common bean weevil, Bruchus obtectus, exhibits a wide
variation in the duration of its comparatively brief death-feint.
Out of 283 freshly emerged beetles, only 72 could be induced to
feign death. Although dropped from a height of 12 inches, hav-
ing their thoraxes pinched, being rolled between the thumb and
forefinger, 211 of the weevils refused to perform in spite of all
the handling. The 72 that actually went into death-feints, fre-
quently did so, with comparatively little trouble. A slight dis-
turbance such as touching them with the tip of a pencil, or causing
them to fall one inch, or gently pressing the sides of the thorax
brought about the desired reaction.

The temperature during these tests was 70° F. For the 72
beetles that reacted, the duration of the feint varied in length
from 1 to 300 seconds. The duration of the weighted, arithmetic
average death-feint was 33.5 seconds, and the standard deviation
was 74.8. The following table shows the duration of the death-
feint in the 72 different beetles that reacted :

No.

beetles

Length of
death-
feint

No.

beetles

Length of
death-
feint

No.

beetles

Length of
death-
feint

No.

beetles

Length of
death-
feint

Seconds

100

105

250

290

300

Harry B. Weiss.

Sept., 1944]

Michener: Culex

263

DIFFERENTIATION OF FEMALES OF CERTAIN
SPECIES OF CULEX BY THE
CIBARIAL ARMATURE

By First Lieutenant Charles D. Michener1
Sanitary Corps, Army of the United States

The purpose of this paper is to describe characters of the
cibarial (or ‘ ‘pharyngeal”) armature of the Culex found in the
southeastern United States. These characters support the usual
subgeneric classification and serve in addition to distinguish
females, previously considered indistinguishable, of certain of
the species commonly placed in the subgenus Melanoconion (e.g.,
by King, Bradley, McNeel, 1942).

The cibarial armature has been used by a number of authors
in separating the subgenera and certain Old World species of
Anopheles. Christophers (1933) gives a good account of the
structures involved in Anopheles.

The armature lies on the posterior margin of the anterior pump
of alimentary canal. This pump has usually been called the
pharynx, but Snodgrass (1943) has pointed out that it is in
reality the cibarium. The second pump is the true pharynx.
The ventral surface of the cibarium is sclerotic. This sclerotized
area ends posteriorly in a broadly concave margin lying between
the two cibarial cornua, which are apodemal muscle attachments
at the posterior end of the cibarium. The cibarial armature,
which is present only in females, consists of a series of sclerotized
projections or teeth along the margin between the cornua.

With a little practice these structures can be studied about as
easily as the male genitalia. The technique used is as follows:
The head is placed in 10 per cent potassium hydroxide and heated
for a short time. Then it is placed in water on a slide and under
a binocular microscope the entire outer wall of the head is broken
away in large pieces by means of fine needles. The pharynx and
cibarium, attached to the hypopharynx, may now be seen within

1 Acknowledgment for both helpful advice and specimens for dissection is
made to Major Stanley J. Carpenter and Captain Woodrow W. Middlekauff.

264

Journal New York Entomological Society

[Vol. LII

and are transferred, with or without separation from the other
mouthparts, to a drop of chloral hydrate medium (Berlese’s
medium) on a slide. Here the pharynx is pulled away from the
cibarium. The cibarial armature will probably still not be
clearly visible because of the imbricated sclerotized teeth in the
membrane of the dorsal surface of the cibarium between the
cibarial cornua. Therefore, this membrane with its imbricated
teeth is dissected away from the dorsal surface of the cibarium in
order to expose clearly the armature on the ventral surface. This
membrane should be preserved, however, since its imbricated
teeth offer characters of importance. The mount is completed
with a coverglass. The separation of individual teeth as recom-
mended by Christophers (1933) for Anopheles has not proved
to be possible with Culex.

The following key separates the species of Culex found in the
southeastern United States into groups on the basis of female
cibarial armatures. Culex atratus Theobald and C. ~bahamensis
Dyar and Knab, found in this area only on the Florida Keys,
have been omitted because of the lack of specimens for study.

1. Cibarial armature consisting of 25 to 35 small, slender teeth; imbricated

teeth of the membrane of dorsal surface of cibarium between cornua

small, lightly sclerotized, brownish 2

-. Cibarial armature consisting of 3 to 8 large, blunt teeth; imbricated teeth
of membrane of dorsal surface of cibarium between cornua large,
heavily sclerotic, blackish (Subgenus Melanoconion ) 3

2. Cibarial teeth not longer than width of a cibarial cornu.

(Subgenus Culex s. str.) pipiens Linnaeus, quinquef asciatus Say,
tarsalis Coquillett, salinarius Coquillett, nigripalpus Theobald
-. Cibarial teeth much longer than width of a cibarial cornu.

(Subgenus Neoculex) apicalis Adams

3. Cibarium with three teeth pilosus (Dyar and Knab)

-. Cibarium with seven or eight teeth.

erraticus Dyar and Knab, peccator Dyar and Knab

These characters are shown in greater detail in the figures.
Except to strengthen the recognized subgeneric classification, the
cibarial characters contribute little to our knowledge of the sub-
genera Culex s. str. and Neoculex.

The three remaining species, however, are externally indis-
tinguishable in the females except that most specimens of errati-

Sept., 1944]

Michener: Culex

265

cus have some coarse, golden, mesoscntal scales and the broad,
appressed, occipital scales are limited to a band along the eye
margin, while in the other two species the mesoscntal scales are
usually dark and the occiput is usually mostly covered with
broad, appressed scales. These characters do not hold for every
specimen. The cibarial characters thus make possible the identi-
fication of female pilosus.

It is interesting to note that Edwards (1932), on the basis of
larval characters, placed pilosus in the subgenus Mochlostryax,

Fig. 1. Cibarial armatures of: 1, Culex {Culex) restuans; 2, Culex {Neo-
culex) apicalis Adams; 3, Culex ( Melanoconion ) pilosus (Dyar and Knab) ;
4 and 5, Culex ( Melanoconion ) erraticus Dyar and Knab.

and erraticus and peccator in Melanoconion. The cibarial arma-
tures of the three species here discussed support this classifica-
tion, but until these characters have been studied for the numer-
ous tropical species of this group, it is not advisable to consider
the cibarial armature as a subgeneric character.

The cibarial armatures of about twenty specimens of C. pilosus
have been examined and found quite uniform in structure. Sev-
eral of those studied were from a series reared from pilosus larvae
by Mr. Wm. V. Reed. The armatures of forty other female

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specimens of Melanoconion were examined. All were seven- or
eight-toothed. Most of them, on the basis of the characters of
the vestiture already mentioned, were presumed to be C. erraticus
and two were from a series reared from erraticus larvas by Lt.
Basil G. Markos. Among the six specimens studied having seven
or eight cibarial teeth but vestiture as in pilosus and peccator,
two from different localities were collected with males of peccator
and are, no doubt, females of that species. Both of these had
but seven teeth.

LITERATURE CITED

Christophers, S. R. 1933. The fauna of British India, Diptera, Vol. IV,
Family Culicidae, Tribe Anophelini, vi + 371 pp., illus.

Edwards, F. W. 1932. Diptera, Fam. Culicidas, in Wytsmann’s Genera
Insectorum, fasc. 194, 258 pp., illus.

King, W. V., G. IT. Bradley, and T. E. McNeel. 1942. The mosquitoes of
the southeastern states, U. S. Dept. Agr., Misc. Publ. 336, 96 pp., illus.
Snodgrass, R. E. 1943. The feeding apparatus of biting and disease carry-
ing flies: a wartime contribution to medical entomology, Smithsonian
Misc. Colls., vol. 104, no. 1, pp. 1-51, illus.

Sept., 1944]

Weiss: Insect Behavior

267

INSECT RESPONSES TO COLORS

By Harry B. Weiss

The purpose of this article is to call attention to the approxi-
mately similar qualitative results obtained by various workers
who used widely different methods of approach and technique in
studying the behavior of insects to different wave lengths of light.
No attempt will be made here to describe the techniques, as they
are already matters of record. In fact they differ as widely as
the methods of approach.

Bertholf (1, 2) exposed the honey bee, and the fruit fly
Drosophila to two translucent glass plates of equal size, one
illuminated with white light and the other with monochromatic
light obtained by means of a quartz prism. The intensity of the
white light was changed until its effect on the photopositive re-
sponse of the insects was equal to that of monochromatic wave
lengths in different portions of the spectrum. He found that for
the honey bee, the stimulating efficiency increased from zero at
6450 A to a maximum of 100 arbitrary units at 5500 A, then
decreased to 10 at 4350 A after which it rose rapidly to a second
maximum of 450 at 3650 A and then rapidly declined to zero
at about 2800 A.

As for Drosophila, Bertholf (2) stated . . starting with
the longer wave lengths the efficiency is very low until it starts
to rise at about 5750 A; from here it rises to a maximum in the
so-called visible spectrum at 4870 A . . . ; from this wave
length it decreases again at 4250 A ; then it rises suddenly and
attains a maximum value at 3650 A . . . ; from here it de-
creases rapidly to zero at 2540 A. ’ ’ Bertholf worked with 30
wave lengths from 2300 A to 7000 A.

Weiss et al. (6, 7, 8) exposed approximately 15,000 insects,
both adult and larval forms, mostly diurnal, but some nocturnal,
involving 40 species in various orders to 10 wave-length bands of
light of equal physical intensities from 3650 A to 7400 A. Each
test lasted from 15 to 30 minutes and a group of 100 or more of
each species was used for from one to three successive tests. The

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Journal New York Entomological Society

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composite group behavior pattern, both for larvas and adult
insects indicated that the stimulating efficiency increased only
slightly from zero at 7200 A to 5750 A. From here it rose to a
maximum at 4920 A. It then declined to a comparatively low
level at 4640 A from which point it ascended to its peak maxi-
mum level at 3650 A.

These authors found that, regardless of the relative positions
of the wave-length bands, the insects made approximately the
same selections time after time, also that when a second and third
test followed the first, there was a shifting of individuals that
went to the different colors, but no change in the final result.
In addition some species such as the Japanese beetle and the Colo-
rado potato beetle responded to what were unattractive wave
lengths under equalized physical intensities, when the intensities
of such unattractive wave lengths were increased. In other
words, it was possible to vary the behavior pattern by varying
the intensities.

Crescitelli and Jahn (3), approached the problem from the
standpoint of the electrical responses of the dark-adapted grass-
hopper eye. “Leads were taken with silver-silver chloride elec-
trodes from fluid-filled chambers about each eye. The entire
surface of one eye was illuminated, and the other eye was kept
in darkness. Records were obtained by means of a cathode ray
oscillograph. For the experiments on colored light Corning color
filters were placed between the light source and the eye.” Six
wave-length bands were employed, extending from about 4000 A
to 7000 A. “The relative intensity transmitted through each
of these six filter combinations was determined by means of a
thermopile and galvanometer. The infra-red radiations were
completely removed from the stimulating light by using 5 cm.
of water and a Corning (AKLO) heat absorbing filter.”

These authors studied the change in form of the electrograms
of the grasshopper eye under variations in intensity of the stimu-
lating light and also the quantitative aspects of the response in
relation to the quality of the stimulating light. They found that
there was apparently no specific effect of wave length on the
electrical response of the whole dark-adapted grasshopper eye.
At equalized intensities there were decided differences in wave

Sept., 1944]

Weiss: Insect Behavior

269

form with the six different spectral bands, but these disappeared
and the color responses were exactly matched when the intensities
of the different spectral regions were properly adjusted. Quoting
again from their paper : ‘ ‘ The form of the electrical response of
the dark-adapted grasshopper eye to brief stimulation by white
or colored light varies according to the intensity of the light.
At very low intensities the response is diphasic, the initial posi-
tive phase of which resembles the a- wave of the vertebrate electro-
retinogram. As the intensity is increased the positive phase
decreases and changes its position while the negative phase be-
comes increasingly prominent. Eventually the positive phase
is completely eliminated and the electrogram takes the form of
the typical high-intensity response. The order of effectiveness
of the different colors in causing this change in wave form is :
green, blue, violet, orange-red, red.”

The curve relating the magnitude of the potential to the wave
length had a peak in the green region of the spectrum, and de-
clined sharply toward the red and less sharply toward the violet.
The magnitude of the electrical response was found to be defi-
nitely related to the quality of the stimulating light and the form
of the response to be influenced by the intensity of the stimu-
lating light, either white or colored.

Jahn and Crescitelli (5), also studied, in the same manner, the
electrical responses of the compound eye of the moth Samia
cecropia, in relation to the quality and intensity of the stimu-
lating light. Part of their conclusions are quoted as follows :
“The electrical responses of the moth and grasshopper eyes to
wave length are surprisingly similar. For both animals the same
type of graph is obtained when the relative magnitude of the
potential is plotted against wave length. This graph has a gen-
eral similarity to the absorption curve of visual purple. Another
aspect of the electrical response to wave length concerns the fact
that no specific effects of wave length on the electrograms are
discernible. By properly adjusting the intensity, the responses
to one color may be exactly matched with the response to any
other color, indicating that the differences in the responses to
different colors of equal intensity are caused merely by differ-
ences in sensitivity and are not effects of wave length per se.”

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In the case of the moth eye the maximum response was obtained
with the green band. The responses dropped sharply toward the
red band and less sharply toward the violet.

Graham and Hartline (4) studying the responses of single
visual sense cells of Limulus to visible light of different wave
lengths found that when the energy of the stimulating light of
different wave lengths was approximately equal, the response to
green was stronger than the responses to either violet or red.
When the energy was increased in the red and violet their level
of response was raised and when the intensities of the different
wave lengths were adjusted so that the responses were equal,
there was no effect of wave length as such, indicating that single
sense cells can gauge brightness, but cannot distinguish wave
length. The relative energies of the various wave lengths re-
quired to produce the same response, after being adjusted in
inverse ratio to the degree of their absorption yielded a visibility
curve for a single visual sense cell that had its maximum in the
green near 5200 A and that declined symmetrically on each side
to low values in the violet near 4400 A and in the red near
6400 A.

Thus the visibility curves of a single visual sense cell of Limu-
lus, although not an insect, of the eye of a grasshopper, a diurnal
insect, and of the eye of a Cecropia moth, a nocturnal insect,
are qualitatively similar to the curve of the relative stimulating
efficiency of different wave lengths of light for Drosophila, as
reported by Bertholf and to the behavior curves for the numerous
adult and larval forms of diurnal and some nocturnal insects as
reported by Weiss et al. These curves are not identical because
of the different methods of approach and technique but they are
all strikingly similar for the visible portion of the spectrum. All
were obtained under wave lengths of equalized physical intensi-
ties. Hartline and Graham and Crescitelli and Jahn by properly
adjusting the intensity were able to match the response to one
color with the response to any other color and Weiss et al., in their
behavior studies found that insects responded to what were un-
attractive colors under equalized intensities, when the intensities
of these colors were increased.

Crescitelli and Jahn (3) report that other authors who worked
with pigeon eyes and the eyes of certain vertebrates also found

Sept., 1944]

Weiss: Insect Behavior

271

that wave form difference are simply intensity differences and
that the electrical response to different wave bands could be
duplicated by adjusting the intensity of the different bands.

Thus it appears that both the electrical responses of the insect
eye and the motor responses of the insect itself to different colors
of equal intensity are due to differences in sensitivity, or to the
absorption of light, which varies with wave length, by the pri-
mary photosensitive substance of the visual sense cells, and are
not the effects of wave length by itself.

REFERENCES

1. Bertholf, L. M. The distribution of stimulative efficiency in the ultra-

violet spectrum for the honey bee. Jour. Agr. Res. 43 (8) : 703-
713. 1931.

2. Bertholf, L. M. The extent of the spectrum for Drosophila and the

distribution of the stimulative efficiency in it. Zeitsch. Yergl.
Physiol. 18: 32-64. 1932.

3. Crescitelli, Frederick, and Theodore Louis Jahn. The electrical

response of the dark-adapted grasshopper eye to various intensities
of illumination and to different qualities of light. Jour. Cellular
and Comp. Physiol. 13 : 105-112. 1939.

4. Graham, C. H., and H. K. Hartline. The response of single visual

sense cells to lights of different wave lengths. Jour. Gen. Physiol.
18 (6) : 917-931. 1935.

5. Jahn, Theodore Louis, and Frederick Crescitelli. ' The electrical

responses of the Cecropia moth eye. Jour. Cellular and Comp.
Physiol. 13 : 113-119. 1939.

6. Weiss, Harry B., Frank A. Soraci, and E. E. McCoy, Jr. Insect be-

havior to various wave lengths of light. Jour. N. Y. Ent. Soc.
49: 1-20; 149-159. 1941. 50; 1-35. 1942. 51:117-131. 1943.

7. Weiss, Harry B., E. E. McCoy, Jr., and William M. Boyd. Group

motor responses of adult and larval forms of insects to different
wave lengths of light. Jour. N. Y. Ent. Soc. 52: 27-43. 1944.

8. Weiss, Harry B. The group behavior of 14,000 insects to colors. Ent.

News 54 : 152-156. 1943.

272

Journal New York Entomological Society

[Yol. LII

HIBERNATION OF THE SYRPHID FLY,
LATHYROPHTHALMUS iENEUS SCOP.

Lathyrophthalmus ceneus Scopoli has the interesting habit of
hibernating gregariously in old nests and burrows of Aculeate
Hymenoptera. On several occasions in the last few winters the
writer has encountered the species in open tunnels of various
anthophorid bees situated in vertical banks or cliffs. In Febru-
ary, 1944, however, several hundred specimens were found hiber-
nating in empty nests of the black and yellow mud-dauber,
Sceliphron servillei (Saussure), in various localities in the San
Joaquin Valley, California. The localities included Merced in
Merced County, Chowchilla and Berenda in Madera County,
Tulare and Tipton in Tulare County, and Kingsburg and Fowler
in Fresno County. In all cases the nests utilized for hibernation
were situated under bridges or culverts. Usually five or six
individuals, more rarely eight or nine, occupied a single empty
mud cell. Prof. F. M. Hull, who very kindly made the identifi-
cation, reported that both sexes were present in a series of ap-
proximately thirty specimens submitted to him, with the females
slightly predominating.

The only other hibernating insect using these nests and ap-
proaching the syrphids in abundance was the elm leaf beetle,
Gallerucella xanthomelcena (Schrank). These, however, were
hibernating elsewhere in the vicinity under a variety of different
conditions, whereas the syrphids appeared to exhibit a marked
preference for this particular habitat. — E. Gorton Linsley.

Sept., 1944]

Clench: Lyc^enid.®

273

TWO NEW SUBSPECIES OF LYC^NOPSIS
PSEUD ARGIOLUS BDV. & LEG. (LEPI-
DOPTERA, LYC/ENIDiE)

By Harry K. Clench
Cambridge, Mass.

The following new subspecies have recently come to my atten-
tion and appear to be worthy of names. As is well known, the
interrelationships of the various named entities of North Ameri-
can Lycamopsis are not as yet satisfactorily determined. The
following two have, therefore, been placed under pseudargiolus
as simple races. It is possible that later on they will be classified
differently.

Lycsenopsis pseudargiolus sidara, new subspecies

Upperside :

Male. Both wings uniform, slightly violet blue, with a thin blackish border
on each. Fringe white, dark toward the apex of the fore wing and occasion-
ally faintly at the vein-ends of both wings.

Female. Both wings shining blue, with a violet tinge. Fore wing with
a broad, dark, marginal border, thickest at the apex. Costa with the blue
whiter and the marginal brown border slightly extended basad. Kind wing:
Costa shaded with brown. Outer margin with a thin black line, basal to
which is a row of spots, one to each interspace. Fringe of both wings similar
to that of the male.

Underside:

Male. Both wings white, grayish in the basal half. Fore wing with the
cell closed by a faint dark line. A thin marginal line runs from costa to
inner margin, basal to which is a row of small spots, each to an interspace.
Basal to these is a scalloped line, sometimes appearing as a row of angled
bars, vertices inward. A post-discal series of heavy short bars runs from
costa to inner margin. The lower (inner marginal) one is very faint, occa-
sionally almost non-existent, but the others are very heavy, and each is set
on the bias, costal end inward, and more heavily so costad than elsewhere.
The costal dash is basally dislocated. Hind wing : The marginal line, series
of dots, scalloped line and cell-end bar as in the fore wing. The post-discal
row of spots is heavy as in the fore wing, but quite dislocated. The two
costal spots are placed inward, the next two outward, and the next four
alternately inward and outward. The base of the wing is scaled with black-
ish. Three basal spots, one on the costa, one in the cell, and one on the

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inner margin, roughly parallel to the body line. These spots, the cell-end
bar and the post-discal series are all outlined faintly in white.

Female. Purer white, lacking the gray basal shading; otherwise similar
to the male.

Holotype, male, Manitou, Colorado, June 5—11, 1882 (ex S. H.
Scudder Coll.).

Allotype, female, same data as holotype.

Paratypes, three males, Starr Ranch, El Paso County, Colorado
(6500 ft.); May 12, 1934; one male, Rock Creek, El Paso Co.,
Colorado (6800 ft.), May 9, 1934; one male, “Col.” (ex S. H.
Scudder coll.).

Holotype, allotype, and four paratypes in the collection of the
Museum of Comparative Zoology. One paratype in the author’s
collection.

Remarks. This subspecies bears a rather close resemblance to
the form named violacea 1 by Edwards. Violacea is represented
in part in the M.C.Z., by a pair of topotypes and a series of speci-
mens labelled “Coalb.,” with various dates around 1883, all re-
ceived through the Scudder collection, they having been received
from Edwards. Sidara may be separated from these specimens
by the closer proximity of the post-discal series of dashes (prin-
cipally on the fore wing below) to the scalloped line. Also, in
sidara the spots of the post-discal series of the hind wing tend to
be more rounded, while in violacea they are either more linear
(usually) or are more quadrate. In sidara the ground color
below tends toward gray, while in violacea it is nearly pure white,
making the white ringing of the discal and basal spots of sidara
absent in violacea. A single male from Cloudcroft, New Mexico,
is apparently of this subspecies, but it is less heavily marked
below, and lacks the basal graying, and the subsequent white-
ringing of the spots.

Lycaenopsis pseudargiolus bakeri, new subspecies

Upperside :

Male. Purplish blue, nearly violet, the costal veins pencilled in whitish
blue. Costa and inner margin of hind wing whitish. Both wings very nar-
rowly bordered by a thin black line. Apical half of the outer margin of the

1 1866, Proc. Ent. Soc., Philadelphia, 6, p. 201. Type loc. : Kanawha River,
West Virginia.

Sept., 1944]

Clench: Lyc^enid^e

275

fore wing narrowly black. Fringe of fore wing white, becoming black to-
wards the apex and at the vein-ends. That of the hind wing white, occa-
sionally faintly dark at the vein-ends.

Female. Fore wing dark brown, with a central area of blue, of varying
extent. A small streak closes the cell. Hind wing also brown, and with a
marginal series of pale blue lunules enclosing small brown spots. Fringe as
in the male.

Underside:

Male. Both wings white. Markings arranged as in other pseudargiolus
forms (see sidara above, for example), the marginal compound border of
both wings rather faint, the post-discal, discal and basal spots very well
marked, dark, and not tending to confluence. Base of hind wing tinged with
greenish.

Female. As in the male.

Holotype, male, Baker, Oregon, April 20, 1941 (J. H. Baker).

Allotype, female, Durkee, Oregon, May 11, 1941 (J. H. Baker).

Paratypes , nos. 1 to 5, male, same data as holotype; nos. 6 to
10, male, Pine Creek, Baker, Oregon, May 26, 1941 ( J. H. Baker) ;
nos. 11, 12, male, Kane Creek, Oregon, March 16, 1934 (F. W.
Lawrence) ; nos. 13, 14, male and female resp., same locality and
collector as allotype, May 19, 1941.

Holotype and allotype in the Museum of Comparative Zoology.
Two male paratypes to be returned to Mr. Baker. Remaining
paratypes in the author’s collection.

Remarks. Both sexes differ from the Californian race echo2 in
the much darker, and therefore more prominent, discal and basal
spots of both wings below. The female above has slightly re-
duced blue areas. Baker i apparently connects, to a greater or
lesser degree, echo with nigrescens Fletcher,3 from which latter
it may be told in the greater amount of blue in the females and
in the constancy of the underside pattern. Quoting Fletcher,
*‘The underside of this variety ( nigrescens ) is remarkable and
specimens of both sexes may be found which, if the underside
alone were seen, might be referred to neglect a, violacea, lucia or
marginata, and some even combine characters of all these. One
beautiful form which frequently occurs, has an irregular, discal,
dark blotch of confluent spots on the secondaries beneath as in

2 1864, Proc. Ent. Soe. Philadelphia, 2, p. 506. Type loc. : California.

3 1903, Trans. Eoy. Soe. Canada, (2) 9, p. 213, fig’d. Type loc.: Kaslo,
B.C.

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[Vol. LII

lucia, and the clear marginal spots of violacea. This form Mr.
Cockle, who has collected this butterfly for several years and has
been much interested in it, considers to be most typical of the
variety.” In all the 16 specimens (14 of them males) of bakeri
examined, there was but one specimen (paratype no. 8) that ex-
hibited the slightest tendency towards the lucia- like confluence
of spots on the underside of the hind wing, and this specimen
was obviously slightly aberrant. The only indication of cliscal
confluence was the basal thickening of the Cui-Cu2, spot of the
post-discal series. The further indication of aberrance was given
by the thickened scalloped line of the submarginal compound
border.

This subspecies is named for Mr. James H. Baker, of Baker,
Oregon, from whom the majority of the specimens were received.

Sept., 1944]

Linsley: Neopasites

277

NEW SPECIES OF NEOPASITES WITH NOTES
CONCERNING OTHERS (HYMEN-
OPTERA, NOMADIDiE)

By E. Gorton Linsley
University of California, Berkeley

Since the publication of a revision of this genus,1 I have had
the privilege of studying additional material through the kind-
ness of Mr. J. N. Knull, Mr. P. H. Timberlake, and Prof. 0. A.
Stevens. The present paper includes the results of this study.

Neopasites elegans Linsley, new species

Female : Form very robust ; color black ; abdomen deep red throughout ;
integument densely, coarsely punctate, moderately dull. Head very densely
punctate, with a large patch of dense appressed white hairs around the
antennal bases, a large white patch behind each eye, the two connected
posteriorly along the posterior margin of the head and the latter by a narrow
median line from the vertex, through the median ocellus to the antennal white
patch; antennas very dark brown, first flagellar segment about as long as
following two together; upper frons and vertex coarsely, contiguously and
subcontiguously punctate, punctures of ocell-ocular area at least as large as
those of disk of mesoscutum ; clypeus densely but more finely punctate ;
labrum clothed with long, dense white hair at base, surface finely punctate,
indistinctly longitudinally carinate for more than two-thirds of its length ;
mandibles black, apices reddish. Thorax coarsely punctate; pronotal collar
densely white pubescent, tubercles margined with white; tegulae very dark
piceous; mesoscutum coarsely densely punctate, margins and anterior three-
fourths of median line, densely clothed with appressed white hairs; meso-
scutellum coarsely, densely punctate, lateral margins and median line densely
white pubescent, the mid-line expanded at posterior margin into a subtri-
angular white patch; metanotum with a white patch on each side; mesepi-
sterna coarsely, very closely, somewhat rugosely punctate, vertical face
broadly margined with dense appressed white hairs; mesosternum with disk
white pubescent. Wings lightly infuscated, veins and stigma brown. Legs
dark brown; anterior and intermediate femora white pubescent beneath at
apex, intermediate and posterior tibiae with posterior face densely, outer face
more thinly, white pubescent. Abdomen deep red, without any dark cloud-
ing; first tergite with a large, suboval white patch on each side bordering

1 Linsley, E. G. 1943. A revision of the genus Neopasites. Trans. Amer.
Ent. Soc., 69: 119-140, fig. 1.

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Journal New York Entomological Society

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lateral margin of basal concavity, the patches separated by nearly twice
their width, basal concavity without white pubescence, tergites two to five
with a large oval white patch on basal margin, the pairs separated by about
their own diameters and becoming successively larger on succeeding seg-
ments except those of the fifth tergite which are smaller, tergites two and
three with a latero-basal white patch on each side those of second tergite
much smaller than the submedian basal patch, those of third tergite scarcely
evident, tergites one to four with a lateral white patch on apical margin,
that of fourth tergite small; fifth sternite bilobate, densely punctate, pubes-
cent, apical margin broadly and shallowly emarginate.

Length 5 mm.

Holotype female (collection of Ohio State University) and one
paratype, female, from El Paso County, Texas, August 30, 1940,
collected by D. J. and J. N. Knu.ll. Two additional paratypes,
female, were taken by Mr. and Mrs. Knull in Culberson County,
Texas, August 30, 1940. One paratype is deposited in the collec-
tion of Mr. P. H. Timberlake, another in the collection of the
writer, the third in the collection of Ohio State University.

In my recent key to Neopasites, this species would run near A.
calliopsidis Linsley but it is larger, more robust, with the integu-
ment duller and more coarsely and densely punctate, the fifth
abdominal sternite more lobate on each side, the disk more densely
punctate and hairy, ‘ and the apical margin broadly, shallowly
emarginate. It further differs in the more extensive white pubes-
cent patches of the head and thorax, the base of the labrum
clothed with white hair, a slightly different pubescent pattern on
the abdominal tergites, and the absence of white pubescence from
the basal abdominal declivity. It is possibly the most beautifully
spotted of the known species.

Neopasites knulli Linsley, new species

Male: Form slender; color black, antennas, mouthparts, legs, and tegulae
brownish or piceous, abdomen red; integument densely, coarsely punctate,
moderately dull. Head very densely punctate, with a large patch of ap-
pressed white hairs around the antennal bases which is denser along the mid-
line where it extends upward nearly to median ocellus, and a large white
patch behind each eye, the two connected posteriorly along posterior margin
of head; antennae brown, distal two-thirds of flagellum reddish, first flagellar
segment about as long as following two together; upper frons and vertex
coarsely, contiguously and subcontiguously punctate, the punctures of ocell-
ocular area a little larger but not quite so dense as those of disk of meso-

Sei>t., 1944]

Linsley: Neopasites

279

scutum; elypeus densely but more finely punctate; labrum with a few long,
erect white hairs at base, surface finely punctate, densely so at base, longi-
tudinally carinate for about one-half its length; mandibles reddish, base and
apex broadly piceous; ventral surface of head shining, area on each side of
gular cavity shining, irregularly punctate, punctures averaging more than
one puncture width apart. Thorax coarsely, very closely punctate; pronotal
collar densely white pubescent, tubercles margined with white; tegulae pice-
ous, margins rufo-testaeeous ; mesose-utum densely, contiguously and more
or less rugosely punctate, anterior and lateral margins and anterior half of
median line clothed with traces of white hairs along lateral and posterior
margins ; metanotum with a white patch on each side ; mesepisterna coarsely,
closely, somewhat rugosely punctate, vertical face broadly margined with
white. Wings very lightly infuscated, veins and stigma light brown. Legs
piceous ; anterior and intermediate femora apically white pubescent beneath ;
anterior and intermediate tibiae thinly, posterior tibiae more densely, white
pubescent externally. Abdomen red; first tergite with a patch of white on
each side bordering lateral margin of basal concavity, the patches separated
by nearly twice their width, basal concavity without white pubescence, ter-
gites two to five with a transverse band of white along basal margin extend-
ing from each side of middle to lateral margin, the pair on tergite two sepa-
rated by about half the distance separating the patches on tergite ohe, those
of each succeeding segment separated by about half the distance separating
those of the preceding tergite, tergites two to four with a small lateral white
patch on apical margin, fifth tergite with an apical fringe of white pubes-
cence, disk faintly clouded with piceous, sixth tergite more distinctly clouded
with piceous; pygidial plate ligulate, at least twice as long as broad; ster-
nites coarsely punctate, apical margins fringed with white pubescence, last
sternite with apical process truncate.

Length 5 mm.

Holotype male (collection of Ohio State University) from Cul-
berson County, Texas, August 30, 1940, collected by D. J. and
J. N. Knull.

This species is superficially very much like Neopasites ( Odon -
topasites) arizonicus Linsley in size, form, and coloration, but
may be distinguished by the naked eyes, less densely punctate
ventral surface of the head, especially on each side of gular
cavity, the short labral carina, and the darker legs and mouth-
parts. It appears to be more closely related to N. (N. ) minimus
Linsley, but the latter species is much smaller (3.5 mm.), with
only a few scattered punctures on the under side of the head,
and the labrum is not carinate.

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Neopasites robertsoni Crawford

Among material recently submitted by Prof. 0. A. Stevens for
study were examples of this species from Hatton, North Dakota,
August 3 (Stevens).

Neopasites stevensi (Crawford)

This species was also taken by Prof. Stevens at Hatton, North
Dakota, on August 3, visiting flowers of Grindelia squarrosa.

Neopasites heliopsis (Robertson)

N. heliopsis was recently recorded1 from Winnecock, North
Dakota. The locality should have read Winnecook, Montana.
The record was based on a specimen in the collection of P. H.
Timberlake.

Neopasites calliopsidis Linsley

This species has been previously recorded from Iowa, Kansas
and Montana. Mr. Timberlake has submitted specimens from
the following localities in Colorado : Boulder County, July 6,
1925 (C. H. Hicks) ; Boulder, June 26 and 28, 1939, on Chry-
sopsis (Timberlake) ; Boulder, 2 miles north, June 25, 1939, on
Chrysopsis (Timberlake) ; and Cuchara Camps, Spanish Peaks,
elevation 8000 ft., July 4, 1939, on Aster (Timberlake). In these
specimens the average extent of the black areas on the abdomen
is greater than in the type series (Iowa), and some males also
have the basal white spots of the tergites confluent on each side.

Sept., 1944]

Weiss: Death-Feints

281

THE DEATH-FEINTS OF ALOBATES PENN-
SYLVANIA DeG., AND ALOBATES
BARBATA ENOCH.

By Harry B. Weiss

Hibernating specimens of both species of darkling beetles were
tested during the last week of April, 1944, in order to determine
the duration of their death-feints. All specimens were kept at a
room temperature of 72° F., several days before testing in order
to fully bring them out of hibernation. Death-feints occurred
when they were picked up or when they were pressed gently,
ventrally or when dropped through a distance of six or twelve
inches. Some beetles required a lot of handling and others very
little in order to bring on the death-feint. The ventral surface
of the thorax appeared to be the most sensitive area. When the
stimulus was applied to the dorsal surface the death-feint did not
occur. It was impossible to apply the stimulus with equal force
each time when it was done by hand, and it is not known if there
is any connection between the force of the stimulus and the dura-
tion of the reaction. However, it does not seem likely that there
is, in view of the fact that a gentle stimulus was just as liable to
promote a long or short death-feint as a hard stimulus. Although
different degrees of pressure and different amounts of handling
were required to initiate the death-feints, both the sensitivity of
the individual and the duration of its reaction probably depend
upon the variable organization of its nervous and motor
mechanism.

Alo~bates pennsylvanica DeG.

Ten specimens of this species were induced to feign death at
Fahrenheit temperatures of 72° and 82°. The durations of the
death-feints are shown in the following table. Some beetles
rested dorsally and others ventrally during the death-feint, but
neither position appeared to affect the duration of the reaction.

Successive death-feints in the same beetle became progressively,
irregularly shorter. One beetle refused to react after the eight-

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Journal New York Entomological Society

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Duration of Death-feint

Beetle

At 72° F. April 19

At 82° F. April 26

Seconds

778

394

266

105

138

262

247

520

230

129

173

1,095

160

620

1,851

380

Average

464.4

268.8

eenth successive death-feint; another after the sixth, and many
of them after the third or fourth.

Alobates barbata Knoch.

The durations of the death-feints of ten specimens of this
species at Fahrenheit temperatures of 72° and 84° are shown
as follows :

Duration of Death-feint

Beetle At 72° F. April 20 At 72° F. April 25 At 84° F. April 25

Seconds

163

165

306

406

180

227

213

414

152

265

340

154

315

475

Average

183.6

179.9

90.0

Successive death-feints in the same beetle of this species were
generally similar to those reported for Alobates pennsylvanica.

Both species reacted alike except for the fact that the death-
feints of A. pennsylvanica endured longer than those of A.
barbata. When the temperature was increased 10 or 12 degrees
the average duration of the reaction declined approximately
one-half, for both species.

Sept., 1944]

Weiss: Death-Feints

283

Apparently there is a variation in the sensitivity of different
individuals, to the external stimuli initiating the death-feint and
also in the period of recovery. These variations are probably due
to differences in the quality of the labile compounds in the
receptive and conductive parts of the nervous system and in the
contractile muscle tissue. The destructive chemical action in-
volved in the reaction to the stimulus and the restoration of the
discomposed substances require different periods of time for dif-
ferent individuals. A high temperature apparently hastens
recovery. It would be of interest to know if internal stimuli
play any part in the restorative processes.

INSECT FOOD HABIT RATIOS OF NEW YORK STATE

By Harry B. Weiss

At various times during the past 15 years I have wondered if
a food habit classification of the species- of insects recorded from
New York would show ratios that differed materially from those
of other sections such as New Jersey, Connecticut, etc. In order
to satisfy my curiosity, I finally classified according to their
family food habits 15,343 of the 15,449 species recorded in “A
List of the Insects of New York,” M. D. Leonard, Editor-in-Chief,
that was published January, 1928, as Memoir 101 of the Cornell
University Agricultural Experiment Station. The Anoplura,
Mallophaga and Siphonaptera were omitted because of their non-
relation to vegetation or to other insects, likewise a few other
species, difficult to classify. Considering the large number of
species involved, these omissions are relatively unimportant.

The difficulty of classifying families of insects in accordance
with the food habits of their members is fully appreciated and
the weaknesses of such a classification have been admitted in a
former paper.1 The terms saprophagous, phytophagous, etc., are
used in their broadest sense and I am aware that such conclusions
as may be drawn from food habit classifications are broad general-
izations.

1 Insect Food Habit Ratios of North Carolina and Mount Desert Island,
Maine. Jour. N. Y. Ent. Soc., vol. 47, p. 155-157, June 1939.

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Journal New York Entomological Society

[Vol. LII

The following table shows the distribution ratios of food habit
types for New York and, for comparative purposes, the ratios for
five other large areas are included. Five of the six areas are
large and embrace a variety of vegetation. Under such condi-
tions one would not expect the distribution ratios of the types of
food habits to vary widely and it will be noted that the New York
ratios are not unlike those for other areas.

No.

Phyto-

Sapro-

Harpacto- Para-

Pollen

feeders,

etc.

species

phagous

phagous sitic

Per cent Per cent Per cent Per cent Per cent

Western Arctic
Coast, N. A.

New Jersey

Connecticut

North Carolina
Mount Desert
Island, Me. ...

New York

400

10,500

6,781

9,249

5,177

15,343

In former papers it was brought out that in relatively small
areas, each with a uniform type of vegetation, the ratios of the
types of food habits, based on the species present, varied in
accordance with the type of vegetation, if the numerical ratios
between the species and the factors tending to reduce their num-
bers are considered as constant. In large areas such as are indi-
cated in the table, the ratios appear to vary but little.

Although the food classifications in the table are broad general-
izations of ‘‘apparent” food, comparatively little has been added
to our knowledge of “actual” food, since B. P. Uvarov called
attention in 1928 (“Insect Nutrition and Metabolism,” Trans.
Ent. Soc. London, Dec. 31, 1928), to the limited number of cases
where the true food of insects was actually known.

Vol. LII No. 4

DECEMBER, 1944

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B, WEISS

Publication Committee

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.

LANCASTER, PA.

NEW YORK, N. Y.

1944

CONTENTS

The Structure of Living Insect Nerves and Nerve Sheaths
as Deduced from the Optical Properties

By A. Glenn Richards, Jr 285

Notes on the Behavior of Burying Beetles (Nicrophorus
spp.)

By Lorus J. Milne and Margery J. Milne 311

Thecla Burdi Kaye, A Synonym

By E. Irving Huntington 328

A New Species of Lambdina and Notes on Two Species
of Besma (Lepidoptera, Geometridae, Ennominae)

By Laurence R. Rupert 329

Psylliidae from Tropical and Semitropical America
(Homoptera)

By John S. Caldwell 335

Notes on Mexican Butterflies, IV

By F. Martin Brown 343

A Revision of the North American Genus Eremomyioides
Malloch (Diptera, Muscidae)

By II. C. I Iuckett 361

Records and Descriptions of Neotropical Crane-Flies
(Tipulidae, Diptera), XVIII

By Charles P. Alexander 369

A Correction 384

The Status and Functions of the International Commis-
sion on Zoological Nomenclature and the Present State
of Its Work

By Karl Jordan 385

Members of the New York Entomological Society 389

NOTICE: Volume LII, Number 3, of the Journal of the

New York Entomological Society was published on

September 11, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.,
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.

Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LII December, 1944 No. 4

THE STRUCTURE OF LIVING INSECT NERVES AND
NERVE SHEATHS AS DEDUCED FROM
THE OPTICAL PROPERTIES* 1

By A. Glenn Richards, Jr.

Zoological Laboratory, University of Pennsylvania

In a previous issue of this Journal relatively indirect histo-
chemical data were presented and were interpreted as indicating
the presence of lipid nerve sheaths in insects (Richards, 1943).
These sheaths, like those of so-called non-myelinated nerve fibers
of vertebrates, are extremely thin and are very labile under the
influence of fixation techniques. Optical studies with polarized
light permit the study of living nerves and so obviate the diffi-
culties attendant upon histological fixation. They also give far
greater sensitivity. The present paper presents data on the
ultrastructure of normal living insect nerves and nerve cords as
deduced from optical data.2 The presence of lipo-protein nerve
sheaths in insects is confirmed and amplified.

The studies reported herein were performed on the nervous
system of adult American cockroaches, Periplaneta americana,
and on larvae of the house mosquito, Culex pipiens. Most of the
studies were made on intact nerve cords and peripheral nerves

2 Thanks are due to Dr. R. S. Bear and Dr. F. O. Schmitt, of the Massa-
chusetts Institute of Technology, for assisting the author with the use and
interpretation of polarized light data, and to Dr. H. B. Steinbach, of Wash-
ington University, for the loan of a Kohler compensator.

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Journal New York Entomological Society

[VOL. LII

but the data so obtained agree with studies on teased, single-fiber
preparations of the larger nerves of cockroaches. Observations
were also made on single specimens of an assortment of other
arthropods to see if similar optical properties occur for nerves
throughout the phylum Arthropoda.

THE USE OF POLARIZED LIGHT

From the viewpoint of optics a nerve is composed of optically
inactive components and several types of optically active com-
ponents. In studies on nerves polarized light is used to deter-
mine (in living or fixed nerves) the presence and status of the
oriented optically active components. One can in a sense frac-
tionate an intact nerve, and with a knowledge of the optical
activities of the extractable components determine where the
various components are located and what their orientation is in
a living nerve. In experimental studies polarized light can be
used to look for degradations of either orientation or chemical
structure. One is dealing, then, with a kind of cytological in-
spection but with a method of inspection that is extremely sensi-
tive when used for nerve studies.

No detailed treatment of the use and interpretation of polarized
light data can be given here. Some of the terms to be used in
this paper may be mentioned in relatively non-technical language
but for an understanding of the complex phenomena involved
reference must be made to the extensive treatises listed below.3

An ordinary beam of light is considered as a series of wave
motions in which the light rays vibrate in all planes 'perpendicular
to the direction of propagation. Plane polarized light is light in
which the vibrations are all in the same plane4 (Fig. 1). As

3 Descriptions of the instruments and general phenomena can be found in
Rogers & Kerr (1933) and other textbooks on mineralogy and optics, espe-
cially Ambronn & Frey (1926). Excellent diagrams of the optical phe-
nomena involved are given by Myers (1938). The most lucid elementary
presentation for beginners is perhaps that of Cheshire (1932). For the
interpretation of biological data see W. J. Schmidt (1937) and Frey-
Wyssling (1938). A more recent and comprehensive analysis of the optical
phenomena shown by nerve sheaths is given by F. O. Schmitt & R. S. Bear
(1939).

4 Actually a light wave consists of electrical and magnetic vectors which
vibrate in mutually perpendicular planes perpendicular to the direction of

Dec., 1944]

Richards: Insect Nerves

287

such, plane polarized light is analogous to parallel waves on water,
where the waves vibrate np and down in one plane while traveling
forward. With polarized light one studies the birefringence
(= double refraction) of a material. This is accomplished by
observing the specimen between crossed Nicol prisms or other

Fig. 1. Diagram of the effect of polarizing blocks set with the polarizing
planes at a 90° angle. The axes of the polarizing blocks is indicated diagram-
matically by parallel lines. The planes in which the light rays vibrate are
sketched in for clarity of three-dimensional visualization.

A light beam traveling along the dotted line in the direction a — > b is
composed of waves vibrating in all axes perpendicular to the direction of
propagation (four of these planes are drawn in this diagram). On passing
through the polarizer A only waves vibrating in the plane of the paper are
transmitted. On reaching the polarizer B, set at a 90° angle, these are just
the waves that are completely extinguished, and therefore no light is trans-
mitted beyond B*

polarizing materials,* * 5 i.e., one prism is below the object and trans-
mits only light that is polarized in one particular plane, the other
prism is above the object and being set at a right angle to the

propagation, but in polarized light studies it is customary to designate the

vibration plane of the electrical vector as the plane of polarization.

5 For a good diagram of how Nicol prisms function see Myers (1938),
figure 84. The same result is obtained by a different principle in ‘ ‘ Polaroid ’ ’
sheets, etc.

* For clarity of diagrammatic presentation the author has taken some
liberties with optics in Figures 1 and 2. Actually, four waves traveling in
phase along one line would combine into a single wave of intermediate posi-
tion and the sum of the four in height. For illustrating the point in question
the method used here seems clearest even though technically different vibra-
tion planes should be drawn along different lines parallel to a-b.

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Journal New York Entomological Society

[Vol. LII

first eliminates all of the light except that which has had its plane
of polarization changed by passing through the specimen (Figs.
1-2). The ability to change the plane of polarization at certain
orientations is a property of birefringent materials. Substances
which show birefringence are said to be anisotropic, substances
which do not are called isotropic. Even with anisotropic sub-
stances, however, there is one direction along which light (ordi-
nary or polarized) is unaltered; this is termed the optic axis of
the substance. The optic axis may be called isotropic, all other
axes are anisotropic. The anisotropic axes have the property of
changing the plane of polarization by splitting any transmitted
beam of light into two rays6 which vibrate in mutually perpen-
dicular planes (Fig. 2) and travel with different velocities.7
The optical effects can be completely described diagrammatically
in terms of a direction of vibration for faster (or slower) trans-
mission (see below) and two refractive indices, since the refrac-
tive index is an inverse statement of the relative speed of light.
Substances with only one refractive index are called isotropic,
substances with two different refractive indices are called uni-
axially anisotropic, and substances with different refractive in-
dices for all three vibration planes of rays perpendicular to the
optic axis are called biaxially anisotropic. Most biological aniso-
tropic substances, including those dealt with in this paper, are
uniaxial.

Several kinds of birefringence are shown by biological mate-
rials. In addition to uniaxial and biaxial anisotropy, birefrin-
gence may be either positive or negative with respect to a particu-
lar direction. These are arbitrary terms applied to differentiate
between materials in which the index of refraction is greater

6 Commonly called the ordinary and extraordinary rays.

7 As Cheshire (1932) points out, birefringent substances include fibrous
and crystalline structures where the orientations of the components are such
that the structure is different in the longitudinal and transverse directions.
In a certain sense the structure is analogous to the grain of wood, and, as
Tyndall showed, the speed of heat and sound waves is different for waves
traveling along the long axis of a piece of wood and for waves traveling
perpendicular to this. With birefringent materials and polarized light we
have similar differences in speeds in different directions with the additional
complication that both the ray direction and the vibration direction are con-
cerned.

Dec., 1944]

Richards: Insect Nerves

289

for the vibration in the stated direction than for the other vibra-
tion perpendicular to it ( positive birefringence) , and materials
in which the relative velocities are the reverse ( negative birefrin-
gence). The direction for faster (or slower) vibration is deter-
mined by comparing the effect with a known standard. Com-
monly one takes advantage of interference effects for this purpose
and uses a gypsum plate which makes the entire field appear red.
Any effect of the specimen must be either additive or subtractive,
and reference to Newton’s series of colors shows which it is.
Knowing the orientation of the various parts of the system one

Fig. 2. Same as Figure 1 but with a sheet of birefringent material (the
specimen, C ) inserted between the polarizers with its axis at a 45° angle to
each. The description is the same as for Figure 1 except that the specimen,
C, oriented at a 45° angle splits the plane polarized beam into two mutually
perpendicular waves vibrating at a 45° angle to the beam from A to C, and
therefore at a 45° angle to polarizer B. The waves do not reach B at a 90°
angle to the vibration direction, and therefore light is transmitted. Adding
to these figures the parts of a compound microscope will give a crude diagram
of the optics of a polarizing microscope of the orthoscope type.

can usually tell by clear-cut color changes (from red to blue or
yellow) whether the substance is positively or negatively bire-
fringent.

The terms positive and negative birefringence, however, can
be used in either of two ways: 1) When the orientation of the
optic axes of the anisotropic substance is known, then the material
may itself be called either positive or negative depending on
whether the index for vibrations parallel to the optic axis is
greater or less, respectively, than the index for vibrations per-
pendicular to this axis. 2) Whether the orientation of the optic

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Journal New York Entomological Society

[Vol. LII

axes is known or not, one can refer to an anatomical structure as
positive or negative with respect to some distinguishing direction.
In many cases in biology the optic axis of the oriented birefrin-
gent material coincides with the long axis of the gross structure
(e.g., muscle, tendon, axis cylinder of nerves, etc.). Thus a
muscle fiber is positively uniaxial in relation to both its long axis
and the optic axes of the anisotropic micelles. In the case of the
nerve sheath, however, the anisotropic micelles are oriented with
their optic axes radial (perpendicular to the long axis of the
nerve). Accordingly, the nerve sheath can be called either
positive or negative depending on whether the reference direction
is the optic axes of the anisotropic particles or the long axis of
the nerve. This can be confusing unless one is careful to note
the reference direction, especially as whichever reference direc-
tion is chosen some nerve sheaths are positive and some negative
in relation to it. In biological literature reference has usually
been made to the anatomically long dimension, and this custom
is followed in the present paper; however in the recent review
by Schmitt & Bear (1939) these authors have shifted to employ-
ing the optic axes as reference directions.

Birefringence may be produced by either the internal structure
of a molecule or its asymmetrical shape. The first is called
intrinsic birefringence, the second is called form birefringence.
Distinguishing between these two types is quite useful in analyz-
ing the source of the observed birefringence. Fortunately this is
readily accomplished. Form birefringence is due to the presence
of oriented submicroscopic rodlets or plates called micelles • im-
mersion of these micelles in a medium of the same refractive
index abolishes their birefringence but has no effect on any in-
trinsic birefringence that may be present. By immersion in
media of various refractive indices one can then determine not
only whether the material possesses any intrinsic birefringence
but more importantly whether there is present a second birefrin-
gent material that is normally masked by the form birefringence
of the first material. Thus, in a so-called non-myelinated nerve
one first observes a birefringence that is positive in relation to
the long axis of the nerve. If such a nerve is immersed in a salt
solution containing glycerine, the sign of birefringence is re-
versed. But if the nerve is first extracted with alcohol the sign

Dec., 1944]

Richards: Insect Nerves

291

cannot be reversed. This is interpreted as due to the positive
form birefringence of the oriented protein normally masking the
negative intrinsic birefringence of the lipids (Schmitt & Bear,
1937).

In treating birefringence quantitatively, certain values can be
measured and others calculated from these measurements. Prop-
erly oriented anisotropic substances split a beam into two rays
which, vibrating in different planes, travel through the specimen
at different velocities. Obviously, if one of the component rays
travels more slowly, it will lag behind and emerge from the
specimen a certain finite distance behind the faster ray.8 This
difference is called the retardation (= phase difference, = ampli-
tude, 4>). With a Kohler compensator the retardation is calcu-
lated from the angle between the compensator plate and the
specimen necessary to compensate for (abolish) the retardation
of the specimen :

where m is the known maximum retardation of the compensator
plate, A the wave length of light (551 mq is taken as the “center
of gravity” of white light), and 3i and (32 the measured angles
using the Kohler method and the Bear & Schmitt modification
respectively (Bear & Schmitt, 1936b). The sign of <p will depend
on whether 3i and j32 are additive or subtractive angles and will
agree with the qualitative determination of the birefringence as
positive or negative in character.

Obviously, the amount the slower ray is behind the faster ray
will depend on both the difference in velocity and the thickness
of the specimen being measured. The difference between the two
velocities, called the magnitude of birefringence, is the funda-
mental property we are interested in obtaining. With sheets of
known composition and orientation the magnitude of birefrin-
gence is readily calculated from :

8 This actually transforms the plane polarized beam of light into an ellip-
tically or circularly polarized beam. However, for our purposes we can con-
tinue to treat it as though we were dealing with two plane polarized beams
vibrating perpendicular to one another.

<p = - sin 2(3i, or
<J> = - 2mA sin 232,

(1)

(2)

(3)

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Journal New York Entomological Society

[Vol. LII

where nt and n2 are the two refractive indices, <p the retardation,
and d the thickness of the specimen.

In nerve studies it is relatively easy to determine the retarda-
tion (<j>) but sometimes difficult to obtain the true effective dis-
tance ( d ). This is partly due to the heterogeneous nature of
nerves, and in the case of nerve sheaths partly to the complica-
tion introduced by the optic axes of the sheath micelles being
oriented radially. Schmitt & Bear (1937) have derived the fol-
lowing equation to allow for the radial orientation of the sheath
micelles :

where n 1 and n2 are the two refractive indices, (32 the compensa-
tion angle determined by the Bear & Schmitt method, dx the axon
diameter or its equivalent, and d2 the axis cylinder diameter or
its equivalent.

For entire insect nerve cords none of these methods for deter-
mining the magnitude of birefringence seems satisfactory. Due
to the low amplitude it has not been found feasible to attempt
quantitative measurements on teased single nerve fibers of in-
sects except for extracted nerve sheaths and the axis cylinder of
stretched nerves. Equation (1) has been used for the points
plotted in Figure 39 but equations (3) and (4) have been applied
to estimate the more fundamental birefringence values for single
fiber preparations and for the neural lamella surrounding the
nerve cord.

For the work reported in this paper a Leitz polarizing micro-
scope was used, employing orthoscopic methods, white light,
various magnifications and several types of compensators. Quan-
titative measurements were made with a Kohler A/20 rotating
mica plate compensator (Leitz Wetzlar No. 2335) using both the
Kohler and the Bear & Schmitt methods (Bear & Schmitt, 1936b).

THE OPTICAL EFFECTS SHOWN BY INSECT NERVES
1. General

Until recently insect nerves have been considered non-myelin-

9 So far as graphs such as figure 3 are concerned, the same curve would be
obtained from using the angle ((3), the retardation (0) and the magnitude
of birefringence ( nx-n2 ).

9.46 sin 2(32

Dec., 1944]

Richards: Insect Nerves

293

ated (Richards, 1943). In many insects ( e.g ., mosqnito larvae)
single nerves are extremely small (less than 2 q in diameter)
but in the large cockroaches some single fibers attain a diameter
of 10 (j and a few are 20-30 q in diameter. The thicknesses of
the nerve sheaths are apparently not more than a few per cent
of the fiber diameter. One can recognize the sheath and estimate
its thickness around single nerve fibers of the cockroach, but the
sheath around nerves of mosquito larvae is beyond resolution with
ordinary microscopical methods.

It is relatively easy to tease out and identify for study single
fibers from a cockroach nerve cord. Many observations can be
made satisfactorily, however, on intact nerve cords containing
many nerves in one bundle. The intact interganglionic abdomi-
nal connectives in the adult American cockroach have a diameter
of 175-280 jj, while those of fourth (last) instar mosquito larvae
are only 13-24 q in diameter.

Around the outside of the intact nerve cord there is a secreted
sheet of material which Scharrer (1939) has termed the neural
lamella. This sheet is also birefringent but does not intefere
with optical analyses of the included nerves. It will be discussed
in a separate section.

In analyzing nerves, they are first examined in a salt solution,
with or without previous fixation, then they may be examined in
glycerine or other media of high refractive index to mask the
form birefringence and so determine what intrinsic birefringence
is present. In insect nerves such immersion normally results in
a reversal of the sign of birefringence, indicating that in a normal
nerve we are dealing with a balance between birefringent mate-
rials of opposite sign (curve 2, Fig. 3). The lipids may be re-
moved by appropriate solvents {e.g., alcohol) ; comparison of the
optical properties before and after extraction permit both show-
ing and localizing the presence of the optically active lipids.
Since the form birefringence of the proteins normally predomi-
nates in non-myelinated nerves, it is ordinarily sufficient to ex-
amine a preparation in salt solution and then in a dilute solution
of glycerine in saline solution. For some special purposes and
for evaluating the method other media, solvents and variations
of technique may be used.

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2. Analysis of the Birefringent Properties of the
Axis Cylinder

Intact, living ventral nerve cords and larger peripheral nerves
of American cockroach adults and mosquito larvae immersed in
saline solutions10 show a slight birefringence which is usually
positive in relation to the long axis of the nerve fibers (Fig. 14) d1
The birefringence of the axis cylinder12 of living cockroach
nerves in salt solution or of fixed cockroach nerves after lipid
extraction is relatively diffuse, is positive in relation to the nerve
axis and seems to be fairly homogeneous throughout single giant
nerve fibers. In the normal relaxed condition the amplitude
of birefringence (4)) due to the axis cylinder is very small but
it can be increased greatly by tension. Cockroach nerve cords
fixed in formol-saline solutions in a relaxed condition, then teased
to give single-fiber preparations, extracted with alcohol (to re-
move masking lipids) and examined in water, show bright posi-
tively birefringent lines for the sheaths but only faint positive
birefringence for the axis cylinder (Fig. 5). Similar results
are obtained for the axis cylinder with living cockroach nerve
cords teased and examined in salt solutions. In contrast, cock-
roach nerve cords fixed in formol-saline solutions in a stretched
condition (1-J times the relaxed length) and then teased and
treated as above show strong positive birefringence throughout
single fiber preparations (Figs. 8-10). Measurements of the am-
plitude of birefringence show that the birefringence of the
sheaths is affected relatively little (perhaps not at all) by the
stretching but that the birefringence of the axis cylinder is
greatly increased. Immersion in glycerine practically abolishes
the birefringence of the axis cylinder showing that this is due
mainly tb anisodiametric micelles exhibiting form birefringence.

10 For cockroach adults the salt solution used contained NaCl 10.93, KC1
1.57, CaCl2 0.85, MgCl2 0.17 and NaHC03 0.17 grams per liter; for mosquito
larvae NaCl 7.8, KC1 0.62, CaCl2 0.40 and NaHCOs 0.17 grams per liter.

11 Occasionally a normal relaxed nerve cord is found to be negative in
saline. Such nerve cords can be reversed to positive by stretching. Natu-
rally, immersion in glycerine-saline does not reverse the sign of birefringence
of such a nerve cord; it only increases the negativity. See curve 3, Figure 3.

12 The axis cylinder is the nerve axon exclusive of its surrounding mem-
brane and sheath.

Dec., 1944]

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Whatever the nature of the tension effect may be,13 the fact re-
mains that the axis cylinder of insect nerves contains anisotropic
micelles, presumably protein, with at least a predominant orienta-
tion in the direction of the long axis of the nerve (Fig. 4).

The studies on single-fiber preparations from cockroaches are
in agreement with observations on intact nerve cords and periph-
eral nerves. In intact nerve cords it is easy to measure the total
birefringence but the superposition of many fibers makes it diffi-
cult to determine what components of the birefringence are due
to the axis cylinder and what to the nerve sheaths. In general, a
longitudinally striated appearance to the birefringence of intact
nerve cords can be interpreted as indicating nerve sheath com-
ponents whereas the more homogeneous birefringence seen in
stretched nerve cords can be interpreted as being due to the sum-
mation of sheaths and stretched axis cylinder effects.

An analysis of the form factor involved in this birefringence
is graphed in Figure 3. Since the measurements were all made
with intact nerve cords these curves, of necessity, show a sum-
mation of axis cylinder and sheath effects. Curve 1 shows the
change in amplitude with change in refractive index of the im-
mersion media for relaxed nerve cords from which the lipids have
been removed by extraction with alcohol and ether. This curve
shows that most of the birefringence of lipid-extracted nerves is
due to micellar form, the micelles having a refractive index be-
tween 1.56 and 1.60 when determined in this manner. A small
residue of intrinsic birefringence is probably also present since
the value never falls quite to zero.14 Comparison of these data

13 There are several conceivable explanations of the increase of birefrin-
gence of the axis cylinder on stretching. Mihalik (1937) has made a similar
report for vertebrate nerves. The phenomenon is being studied further. It
is presented here only because the birefringence of the axis cylinder of
insect nerves can be seen most clearly in stretched nerves.

14 This is the interpretation accepted by Schmitt and Bear. The Ambronn
immersion technique commonly encounters difficulties with biological mate-
rials. Theoretically curve 1 should be symmetrical. Correction for the
shrinkage that occurs in media of higher refractive index is not sufficient to
make these curves symmetrical. Such slight asymmetry is seen in various
published curves ( e.g ., Chinn & Schmitt, 1937, Fischer, 1944). Different
methods of fixation give roughly parallel curves of considerably different
height (Bear, Schmitt & Young, 1937b). Curve 1 and 2 of the present paper

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with the data from single-fiber preparations suggests that the
form birefringence plotted in curve 1 is due to both the axis
cylinders and the nerve sheaths but how much is contributed by
each of these components has not yet been determined. Curve 2
shows the change in amplitude of a formalin-fixed nerve cord
immersed in media of increasing refractive index with reversal
of sign at approximately 1.343. Curve 2 is typical for normal
cockroach nerves. Theoretically one would expect this curve to
parallel curve 1 if a series of non-lipid-solvent immersion media
of higher refractive index were available. Curve 3 gives data
from a similar immersion series for one of the occasional nerve
cords which is already negative in saline. Great variation, such
as is shown by the difference between curves 2 and 3, occurs be-
tween different preparations. This seems, at least in part, due to
tension, and will be discussed in a subsequent paper.

It does not seem feasible to attempt estimating the true mag-
nitude of birefringence (nx — n2) from measurements on entire
nerve cords. Estimates from formalin-fixed, alcohol-extracted,
single-fiber preparations of the cockroach using equations (1) and
(3) indicate that the magnitude of birefringence of the axis

and similar curves by Chinn (1938) though roughly parallel are not so far
apart as one would expect. More serious is the great variation of curves for
some materials depending on the immersion series employed. Castle (1936)
has treated in some detail the radically different types of curves shown by
chitin in different series of immersion media where imbibition and possibly
adsorption alter the picture and complicate interpretation. The use of dif-
ferent series of immersion media has given only slight differences for nerve
sheaths (Chinn & Schmitt, 1937) but a much lower value for the refractive
index is assumed from glycerine -immersion series in the recent paper by
Werndle & Taylor (1943). The complex phenomena involved in these unex-
pected variations are not understood. Castle (1936) feels that the situation
is so complex that it defies explanation at present, and that the immersion
method is not valid for the determination of the refractive index of certain
materials such as chitin. The curves from nerves and nerve sheaths by
various methods and media are, however, sufficiently consistent to be reason-
ably certain that '(1) insect nerves are similar to other nerves, and (2) that
the protein component shows largely a form birefringence and that it is of
opposite sign from the lipid birefringence. Under the circumstances it
seems that the best proof of the lipo-protein nature of the nerve sheaths comes
from the results of extraction experiments rather than from the immersion
curves (metatropic effect).

Dec., 1944]

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•cylinder ranges from nearly zero (immeasurably low) in relaxed
nerves to approximately 0.0018 for stretched nerves.12 The sev-
eral possible reasons for low magnitudes in axis cylinders are dis-
cussed by Bear, Schmitt & Young (1937b) and Richards, Stein-
bach & Anderson (1943).

Fig. 3. Data from immersion experiments with cockroach nerve cords.
Entire interganglionic connectives were used; the measurements accordingly
represent the summation of effects shown by a large bundle of nerves. Curve
1 gives the averages from measurements on six nerve cords fixed in 95%
ethyl alcohol and subsequently measured in media ranging from 1.331 to 1.659
in refractive index,* with corrections being made for differences in size and
for shrinkage in some of the media. Curve 2 gives the readings from a
formalin-fixed nerve cord immersed in distilled water and a series of increas-
ing concentrations of glycerine, range 1.333 to 1.435 in refractive index.
This curve closely approximates the typical picture for fresh nerves in saline,
with reversal of sign occurring at a glycerine concentration between 10 and
15%. Curve 3 gives measurements from one of the few formalin-fixed nerves
which was found to be negative in water.

* The immersion media used were: methyl alcohol, ethyl alcohol, 1-nitro-
propane, methylcyclohexane, chloroform, carbon tetrachloride, benzene, di-
methyl phthalate, methyl salicylate, anethole, aniline, bromoform, carbon
bisulfide and alpha-bromo -naphthalene.

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The nerve cords of mosquito larvae are much smaller than those
of cockroaches, and the included nerves are all very small ( < 1 to
2 (j ) . All studies on mosquito nerves were made with intact nerve
cords since satisfactory single-fiber preparations were not ob-
tained. The intact living nerve cord in appropriate salt solution,
like that of the cockroach, is positively birefringent in relation to
its long axis. In the relaxed condition (Fig. 14) it shows a more
or less striated birefringence which, as noted above, is interpreted
as indicating the presence of a relatively strongly birefringent
sheath component. In the stretched condition the birefringence
is stronger and nearly homogeneous. By comparison with cock-
roach nerves, this is interpreted as indicating an increase of the
axis cylinder component of birefringence. The data from mos-
quito nerve cords are consistent with the idea that they have
qualitatively the same configuration as is found in cockroach
nerves, but they are very small and the amplitude of birefrin-
gence is very low.15

There is no reason to think that the ultrastructure of the axis
cylinder of insect nerves is different from that of nerves of other
animals. The amplitude of birefringence of relaxed insect nerves
is unusually low but this may possibly be reconciled by further
study. Otherwise, the picture recorded above agrees well with
reports by various workers on various nerves (see W. J. Schmidt,,
1937). The birefringence of the axis cylinder is due to aniso-
tropic protein micelles showing chiefly a form birefringence
(anisodiametric particles). These micelles are oriented or at
least predominantly oriented with their optic axes parallel to
the long axis of the nerve since the axis cylinder is isotropic in
cross section. Their birefringence is positive both with respect
to their optic axes and to the nerve axis (Fig. 4). As already
reported with other nerves, there is a slight reduction in the
amplitude of birefringence on histological fixation. There is a
rapid decay or loss of birefringence of the axis cylinder follow-

15 The phase retardation (0) of an entire, relaxed, 18 (a, interganglionic
connective of a mosquito larva in saline as calculated from equation (1) is
of the order of only 2-3 mp. In a stretched nerve cord this may rise to
6-8 mp. These values compare favorably with values given for cultured
Corethra nerves by Pfeiffer (1943).

Dec., 1944]

Richards: Insect Nerves

299

ing mechanical injury, the action of certain venoms and insecti-
cides, and attending post-mortem degeneration.16

3. Analysis of the Birefringent Properties of the
Nerve Sheaths

Insect nerve sheaths are relatively thin. In the largest nerves
of the cockroach they attain a thickness of only a little more than
one micron, i.e., they are only a few per cent of the fiber diameter.
In smaller nerves they seem to be proportionately thinner, and in
the smallest cockroach nerves and in all nerves of mosquito larvae
they are too thin for direct observation (submicroscopic thick-
ness) . Due to the thinness of these sheaths it is difficult to deter-
mine for certain whether or not they are produced by distinct
sheath cells. However, in longitudinal sections of insect nerve
cords one can find a few nuclei between the nerve fibers.17 Bear,
Schmitt & Young (1937a) concluded that nuclei found around
squid giant nerve fibers probably represented cells analogous to
the Schwann cells of vertebrate nerves. Possibly the nuclei seen
in insect nerve cords likewise represent sheath cells analogous
to Schwann cells.

These thin nerve sheaths are metatropic, i.e., they exhibit a
weak positive birefringence (in relation to the nerve axis) which
is readily reversed to negative by immersion in media of slightly
higher refractive index (curve 2, Fig. 3). These data can be
interpreted in the same manner that similar data from other ani-
mal groups have been interpreted by Schmitt & Bear (1937,
1939). The individual nerve sheaths are composed of bound
layers of protein and lipid, both of which are birefringent but
of opposite sign. The protein normally dominates slightly with
its form birefringence. Masking the form birefringence of the
protein with glycerine permits the intrinsic lipid birefringence to
be seen. Removal of the lipids with appropriate solvents ( e.g .,
alcohol) increases the positive birefringence and makes it impos-
sible to reverse the sign of birefringence with glycerine. Since
two oppositely birefringent compounds can be thus demonstrated

16 A presentation of some of the degenerative effects due to venoms and
insecticides will be given in subsequent papers.

17 Other than those associated with tracheae.

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and identified (Figs. 5 vs. 11, 14 vs. 15), it follows that normally
both are present in the sheath, and that the normal sheath bire-
fringence is a picture of how much one component (in this case
protein) predominates in birefringence over the other.

The axis cylinder viewed from the side is birefringent through-
out. The nerve sheaths, however, show birefringence only at or
very near to the edges of the nerve. This is due to the manner
of orientation of the optic axes of the anisotropic micelles. In all
birefringent materials and structures there is' one axis, the optic

Fig. 4. A diagrammatic representation of the orientation of the birefrin-
gent micelles in a nerve and its sheath. The long axes of the rodlets in
these diagrams is used to indicate the orientation of the optic axes of the
micelles. On the left is a hemi-section of a nerve showing the micelles of
the axis cylinder oriented longitudinally while those of the sheath are
oriented radially. On the right is an enlarged sketch showing the arrange-
ment of bound lipid and protein micelles in concentric layers as suggested by
Schmitt, Bear & Palmer (1941).

axis, along which the structure is isotropic. In the nerve sheaths
both the protein and lipid molecules must be oriented with their
optic axes arranged radially (Fig. 4). With such an arrange-
ment, the central part of the sheath should appear isotropic, as
it does, since the optic axes of the micelles lie in the axis of the
microscope, whereas the edges should be strongly birefringent, as
they are. This is confirmed by observation of negative polariza-

Dec., 1944]

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tion crosses observed in cross sections of alcohol-extracted giant
cockroach nerves (compare Chinn & Schmitt, 1937).

The magnitude of birefringence (Wx-Wa) for the sheath can
be calculated approximately from measurements on single giant
fibers of cockroaches. Fresh nerves in saline show only extremely
faint sheath birefringence. This indicates that the balance of
birefringence from the protein and lipid (opposite signs) is
nearly equal, and the magnitude of birefringence of the combina-
tion is therefore nearly zero. Alcohol-extracted nerves (Figs.
5-7) should show only the protein component. The average of
measurements on eleven alcohol-extracted nerves ranging from
6 to 10 p in diameter using equation (4) is the extremely low
value of nt - n2 = 0.0016. This represents the value for sheath
protein ; the lipid value must be similar but slightly lower.

The nerve cords of mosquito larvae show the metatropic reversal
as well as cockroach nerves, but the small size of the fibers makes
it difficult to show positively that the reversal is in the sheath.
Presumably the situation is the same as in cockroaches except for
the small size and low amplitude (Figs. 14—15).

The protein component of the individual nerve sheaths of in-
sects may be collagenous as in other animals (see e.g., Schmitt,
Hall & Jakus, 1942). Direct proof of this was not obtained but
alcohol-extracted single-fiber preparations from cockroaches ob-
served during treatment with dilute acetic acid showed a reduc-
tion and eventual loss of sheath birefringence. This would be
expected for collagenous sheaths.

The lipid component of the individual nerve sheaths of insects
seems almost certainly to include phosphatids as in other animals
(Schmitt, Bear & Palmer, 1941). The solubilities of the sheath
components are consistent with the idea that they include phos-
phatids. Also it is possible to extract considerable quantities of
phosphatids and other myelin constituents from bee brains (un-
published data by Dumm, Patterson & Kichards).

Metatropic nerve sheaths are found on axons throughout the
insect. They are best studied from interganglionic connectives
and large peripheral nerves {e.g., cereal nerve) but they can be
demonstrated throughout ganglia and in small distal peripheral
nerves. Likely these sheaths extend over the nerve cell bodies

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themselves but this was not studied. Chinn (1938) reported
that a sheath continuous with and similar to the axon sheath
extends over the nerve cells of lobsters, crayfish and leeches.

The metatropic nerve sheaths of cockroaches and mosquito
larvae exhibit numerous of the properties already described for
other nerves. Birefringence is quickly lost following mechanical
injury. The metatropic effect is lost on air-drying. Absolute
ethyl alcohol removes the lipid quickly but weak alcohol removes
it slowly with the production of birefringent particles that are
stainable with Sudan dyes (Richards, 1943) . Other lipid solvents
except acetone also quickly block the metatropic effect (phospha-
tids are relatively insoluble in accetone) . The metatropic reversal
is most readily demonstrated by immersion in glycerine solutions
but it can also be produced by other solutions of high refractive
index (e.g., sucrose) and is independent of shrinkage. The
amplitude is somewhat reduced by fixation with formalin, etc.,
but after fixation with non-lipid solvents the lipids are more
strongly bound and difficult to remove (see Mezzino, 1931).
Likely there is also a relation between fiber diameter and ampli-
tude of birefringence ; certainly the larger nerves of cockroaches
show more birefringence than the small nerves (see Schmitt &
Bear, 1939, Fig. 1).

Cockroach nerve cords degenerating in saline show first a
decrease in amplitude and then loss of the birefringence of the
axis cylinder, then a gradual decrease and loss of the lipid com-
ponent of birefringence. Nerve cords degenerating in glycerine-
saline solutions, however, give rise to sphaerites which gradually
decrease in size and eventually disappear (Fig. 16). These bire-
fringent sphaerites originate from the sheath lipids, they can be
produced by weak alcohol or degeneration in glycerine but have
not been found to occur in cockroach nerve cords degenerating
in saline (compare Baldi, 1929). The lipid component can also
be abolished by the action of cobra venom which is known to con-
tain phosphatase enzymes. They are also affected by certain
insecticides ; these data will be treated in subsequent papers.

Insect nerve sheaths differ quantitatively from other nerves
in the low order of birefringence and especially in the very close
balance of positive and negative components. The balance of

Dec., 1944]

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303

these components seems closer in insect nerve sheaths than in
other recorded metatropic nerves. Only 10-15 per cent glycerine
is needed to effect reversal of sign in the insects studied, whereas
30 per cent glycerine is required for Crustacea, 35-40 per cent
for squid giant fibers, 50 per cent for non-myelinated spinal
ganglion cells of the frog, and 70 per cent for leech nerves. In
all of these the refractive index of the anisotropic micelles is
reported to lie in the range 1.56 to 1.60 ; accordingly this varia-
tion in necessary refractive index of the immersion media likely
indicates roughly the ratio of protein birefringence to lipid
birefringence in the various nerves.

METATROPIC NERVE SHEATHS IN OTHER ARTHROPODS

A hasty survey was made covering a number of scattered
arthropods to see if the type of nerve sheath studied in the cock-
roach and mosquito is to be found throughout the Annelid-
Arthropod complex. All species examined exhibited metatropic
nerves. Species examined by the author were an unidentified
spider, a centipede ( Scutigera sp.), a millipede ( Fontaria sp.),
a caterpillar ( Lymantria dispar ), a beetle ( Scarites subterranus)
the honey bee (Apis mellifica) and also a marine polychaete worm
(Nereis virens). Other authors have already reported on the
earthworm (Lumbricus sp.), the medicinal leech (Hirudo medi-
cinalis), Limulus and various Crustacea (prawn, shrimp, crayfish,
crab, lobster).

With all the major groups of the Annelid- Arthropod complex
except the Onychophora represented in the above list, it certainly
seems probable that Schmitt & Bear were correct in suggesting
that metatropic nerve sheaths would be found throughout these
phyla.

THE NEURAL LAMELLA

Around the outside of the nerve cord and of peripheral nerves
is a homogeneous sheet secreted by an underlying layer of non-
nervous cells. Scharrer (1939) studied this rather extensively
in cockroaches and termed the cell layer the “ perineurium’ ’ and
the secreted sheet the 4 * neural lamella.” She pointed out that
the neural lamella is optically homogeneous and stains with dyes
like the connective tissue of vertebrates. It is several microns

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thick in cockroaches but is too thin to measure in mosquito larvae.
It seems to be present in all insects but is commonly very thin
and inconspicuous.

The neural lamella is strongly birefringent. It shows as a
bright line along the edge of all nerve cords and is positively
birefringent in relation to the long axis of the nerve cord (Fig.
12). 18 In cross sections it shows a strong negative polarization
cross. Immersion experiments show that its birefringence is
almost entirely due to form birefringence, but attempts to deter-
mine the refractive index by immersion methods encountered the
same sort of difficulty recorded by Castle (1936) for chitin. Im-
bibition or oriented imbedding in some cases gives quite different
measurements for media of the same refractive index. The re-
fractive index seems most likely in the neighborhood of 1.47, and
clearly is different from both chitin and the sheaths (collagen ?)
around individual nerves. Although the neural lamella shows as
a bright line due to its depth, the true magnitude of its birefrin-
gence is quite low. Using equation (4), nx-n2 is found to be
approximately 0.00022.

The neural lamella is unaffected by extraction with lipid sol-
vents. It is completely and readily dissolved by strong alkali,
and accordingly is not chitinous. It gives a strong protein reac-
tion (xanthoproteic test) but seems not to be collagen since it
does not swell, dissolve or even lose its birefringence in dilute
acetic acid (3 days) and since immersion experiments give differ-
ent results for the neural lamella and the presumably collagenous
sheaths around individual nerves. Serial sections show that the
neural lamella is composed of concentric thin layers. Attempts
to obtain electron micrographs showed only that the neural
lamella fractures in an irregular manner suggesting a non-fibrous
structure. Stretching experiments show that the neural lamella
is elastic and possesses strong, photoelastic properties.

is The neural lamella does not interfere seriously with a study of the
birefringence of the included nerves. It shows only as a bright line along
the edge, and can be ignored. Proof that it does not interfere with readings
made on the included nerve bundle comes from experiments in which the
same reading was obtained for the nerve bundle before and after manual
removal of the neural lamella.

Dec., 1944]

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305

The above data suggest that the neural lamella of cockroaches
is a series of elastic, homogeneous, concentric sheets composed of
anisodiametrie protein micelles exhibiting form birefringence and
arranged with their optic axes perpendicular to the surface of
the sheet. The negatively uniaxial protein micelles are arranged
with their optic axes at right angles to the nerve axis, and accord-
ingly the neural lamella appears positively birefringent in rela-
tion to the axis of the nerve cord.

No attempt has yet been made to study accurately the per-
meability of the neural lamella but obviously it must be per-
meable. Exchanges with the blood must take place through it,
fixing fluids used in histology penetrate rapidly, and even the
rather large aggregates of solubilized detergents — Black Sudan
B (a polyazo dye) penetrate the neural lamella more rapidly than
they will dialyze through a collodion membrane.

The birefringence of the neural lamella of a wasp and a beetle
has already been described and figured by W. J. Schmidt (1937,
p. 273, Fig. 70), but seemingly Schmidt erroneously thought that
this sheet compared with the sheaths of single vertebrate nerves.

DISCUSSION

The data presented in this paper supplement those given pre-
viously (Richards, 1943), and show that the structure of insect
nerves and nerve sheaths is closely similar to the structure re-
ported for other animals (see Schmitt & Bear’s review, 1939).
This structure is diagrammed in Figure 4. Around the outside
of insect nerves and nerve cords is a secreted sheet, the neural
lamella, which while serving the same protective purpose is dif-
ferent from the protective coatings around vertebrate nerves and
nerve cords.

Few observations on the birefringence of insect nerves are to
be found in the literature. Bruno (1931) reported that insect
nerves (various species) are isotropic. This bare statement can-
not be evaluated in the absence of any information on the method
of preparation and the type of compensator employed. Perhaps
the negative report is simply an expression of failure to recognize
the extremely low amplitude of relaxed insect nerves. W. J.
Schmidt (1937) has already refuted Bruno’s report. He pub-

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lished a figure showing the birefringence of a fresh peripheral
nerve from the head of a wasp (Fig. 70, p. 273) and described
similar results obtained with the ventral nerve cord of a beetle.
From his brief comments, however, it is not certain how clearly
he distinguished the various components of the birefringence.
Certainly the sheath to which he refers is the neural lamella ; he
seems not to have seen the sheaths of the individual nerves or to
have determined their lipo-protein character. Recently Pfeiffer
(1943) has studied the growth of single nerves from Corethra
larvse (Diptera) in tissue culture. He records a positive uni-
axial birefringence for normal Corethra nerves, with an ampli-
tude of birefringence comparable to that described herein for
mosquito larvae. Pfeiffer was interested in studying nerve
growth ; he does not mention nerve sheaths and seems not to have
distinguished between sheath and axis cylinder effects or to have
recognized the presence of metatropic sheaths.19

Numerous references can be found in entomological literature
stating that insect nerves are non-myelinated. These statements
are readily understandable in view of the extreme thinness of the
sheaths (commonly submicroscopic) and the correspondingly low
lipid content.

SUMMARY

1. Insect nerves and nerve cords in saline solutions show a
positive uniaxial birefringence in relation to the length of the
fiber. This is reversed to negative by immersion in media of
higher refractive index, i.e., the nerves are metatropic. Qualita-
tively they seem to agree well with the structure of other so-called
non-myelinated nerves as recorded by Schmitt, Bear and others.
A diagram presenting the orientation of the optically active
micelles is given in Figure 4.

2. The axis cylinder of insect nerves shows largely form bire-
fringence which is positive in relation to both the nerve axis and
the optic axes of the micelles. Most of this birefringence is due

T9 The extremely small size of these nerves may have caused Pfeiffer to
overlook the sheath. In nerves of comparable size from mosquito larvse the
sheaths are Of submicroscopic thickness. Another possibility is that insect
nerves growing in tissue culture may possibly differ from nerves growing in
an animal.

Dec., 1944]

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307

to micellar form but a small residue of intrinsic birefringence
seems to be present. The amplitude of birefringence of the axis
cylinder increases greatly on being stretched.

3. Each insect nerve is surrounded by a discrete sheath of lipo-
protein. In thickness these sheaths are at most only a few per
cent of the fiber diameter, and accordingly in the case of small
nerves are too thin to be observed directly (submicroscopic).
This sheath is responsible for the metatropic reversal effect.
Both the lipid and protein components are oriented with the
optic axes of their micelles arranged radially. They are of
opposite sign. Normally the nerve sheaths appear positively
birefringent in relation to the nerve axis because the positive
component due to the proteins slightly overbalances the negative
component due to the lipids. Removal of the intrinsically bire-
fringent lipid increases the positive birefringence and prevents
metatropic reversal. Masking the form birefringence of the
protein permits the intrinsic birefringence of the lipids to be
seen (metatropic reversal).

The protein component of the sheath may be collagen. Its
micelles are negatively birefringent in relation to their optic
axes but being arranged radially make the nerve appear posi-
tively birefringent in relation to its length. Its birefringence
is mostly due to micellar shape but a small residue of intrinsic
birefringence seems to be present.

The lipid component of the sheath is probably a mixture of
phosphatids with perhaps other “myelin” components. The
lipid micelles are positively birefringent in relation to their optic
axes but being arranged radially make the nerve appear nega-
tively birefringent in relation to its length. The lipid birefrin-
gence is intrinsic since it is unaffected by the refractve index of
immersion media.

4. Insect nerves seem to differ from the non-myelinated nerves
of other animals chiefly in the low order of amplitude of birefrin-
gence and the extreme thinness of the nerve sheaths. These
seeming differences may possibly be illusory since ordinarily only
large nerves are studied whereas insect nerves are commonly very
small.

5. Metatropic nerve sheaths have been reported to date in an
earthworm, sandworm and leech (Annelida), a spider and Limit-

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[Vol. LII

lus (Arachnida), various shrimps, crabs, etc. (Crustacea), a
millipede (Diplopoda), a centipede (Chilopoda) and five orders
of pterybote insects (cockroach, mosquito, moth, bee, beetle). It
certainly seems probable that at least for the Annelid- Arthropod
complex Schmitt & Bear were correct in suggesting that meta-
tropic nerve sheaths will be found throughout the invertebrates.

6. Around the nerve cord and peripheral nerves is a secreted
permeable elastic layer, the neural lamella, which is structurally
and chemically different from the protective coatings around verte-
brate nerves. The neural lamella is or at least contains a protein
which shows form birefringence and photoelastic properties.
This protein seems to be different from collagen and different
from the protein of the sheaths around individual nerves. Im-
mersion experiments are complicated by the oriented imbedding
of imbibed media. The neural lamella is composed of thin con-
centric layers with the micelles arranged perpendicular to the
surface (radial in relation to the nerve). These micelles, like
sheath proteins, are negatively uniaxial in relation to their optic
axes; being arranged radially they make the sheet appear as a
positively birefringent line (in relation to the nerve axis) along
the edge of the nerve.

LITERATURE CITED

Baldi, F. 1929. Richerche sulla degenerazione Walleriana. Prime in-
dagini a luce polarizzata.- Riv. Neurol., 2 : 56-63.

Bear, R. S. and F. O. Schmitt. 1936. The optics of nerve myelin. Jour.
Optical Soc. Amer., 26: 206-212.

. 1936. The measurement of small retardations with the polarizing

microscope. Jour. Optical Soc. Amer., 26: 363-364.

Bear, R. S., F. O. Schmitt and J. Z. Young. 1937. The sheath components
of the giant nerve fibers of the squid. Proc. Roy. Soc. London, B,
123: 496-504.

. 1937. The ultrastructure of nerve axoplasm. Ibid., 123 : 505-519.

. 1937. Investigation of the protein constituents of nerve axoplasm.

Ibid., 123 : 520-529.

Bruno, G. 1931. Sulla struttura dei nervi di aleuni insetti. Monit. Zool.
Ital., 42: 8-15.

Castle, E. S. 1936. The double refraction of chitin. Jour. Gen. Phys., 19:
797-805.

Cheshire, F. J. 1932. The polarization of light. Jour. Television Soc.,
ser. 2, 1: 201-207.

Dec., 1944]

Richards: Insect Nerves

309

Chinn, P. 1938. Poralization optical studies of the structure of nerve cells.
Jour. Cell. Comp. Phys., 12 : 1-21.

and F. O. Schmitt. 1937. On the birefringence of nerve sheaths

as studied in cross section. Jour. Cell. Comp. Phys., 9: 289-296.
Fischer, E. 1944. The birefringence of striated and smooth mammalian
muscles. Jour. Cell. Comp. Phys., 23 : 113-130.

Mezzino, L. 1931. Rifiessioni ed osservazioni istologiche sulle fibre nervose
midollate. Riv. Biol., 13 : 31-56.

Mihalik, P. 1934. Mechanisch-experimentelle Untersuchungen fiber die
Doppelbrechung der Markhaltigen Nervenfaser. Zts. Zellforsch.
mikr. Anat., 21: 653-656.

Myers, L. M. 1938. Television Optics. 2nd Ed. 362 pp. Pitman & Sons,
London.

Pfeiffer, H. H. 1943. Polarizationsmikroskopische Messungen an gerichtet
gespannten Mutterstficken von Ganglienkulturen in vitro. Natur-
wiss., 31 : 47-48.

Richards, A. G. Jr. 1943. Lipid nerve sheaths in insects and their probable
relation to insecticide action. Jour. N. Y. Ent. Soc., 51: 55-69.

, H. B. Steinbach and T. F. Anderson. 1943. Electron microscope

studies of squid giant nerve axoplasm. Jour. Cell. Comp. Phys.,
21: 129-143.

Scharrer, B. 1939. The differentiation between neuroglia and connective
tissue sheath in the cockroach (Periplaneta americana). Jour.
Comp. Neurol., 70: 77-88.

Schmidt, W. J. 1937. Die Doppelbrechung von Karyoplasma, Zytoplasma
und Metaplasma. Protoplasma Monographien, 11: 1-388.

Schmitt, F. O. and R. S. Bear. 1937. The optical properties of vertebrate
nerve axons as related to fiber size. J our. Cell. Comp. Phys., 9 :
261-273.

. 1939. The ultrastructure of the nerve axon sheath. Biol. Rev., 14:

27-80.

and K. J. Palmer. 1941. X-ray diffraction studies on the nerve

myelin sheath. Jour. Cell. Comp. Phys., 18: 31-42.

, C. E. Hall and M. A. Jakus. 1942. Electron microscope investi-
gations of the structure of collagen. Jour. Cell. Comp. Phys., 20:
11-33.

Werndle, L. and G. W. Taylor. 1943. Sheath birefringence as related to
fiber size and conduction velocity of catfish Mauthner, Mfiller and
peripheral fibers. Jour. Cell. Comp. Phys., 21: 281-293.

PLATE X

In all figures the cross-hairs indicate the orientation of the Nicol prisms
of the microscope. Extra contrast and compensation obtained with a Kohler
V20 rotating mica plate compensator. Photographs, of course, do not dis-
tinguish between positive and negative birefringence; this is determined by
compensators and stated in text and legends.

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Journal New York Entomological Society

[Vol. LII

Fig. 5. Teased single large nerve fiber from cockroach nerve cord fixed
in 95% ethyl alcohol in relaxed condition. Lipids extracted by alcohol. In
distilled water; birefringence positive in relation to nerve axis. The parallel
light lines represent the protein component (collagen ?) of the nerve sheath.
Diameter of this fiber 16 p,. Magnification 300 X.

Fig. 6. A group of medium-sized nerves from the same preparation as
Figure 5. Diameters of these fibers 5-6 p. Magnification 300 x.

Fig. 7. A single small fiber, 3 p in diameter, from same preparation. Note
how faint the sheath is. Magnification 300 x.

Fig. 8. A single teased cockroach nerve fiber which was fixed in 95%
alcohol while stretched to approximately li x its relaxed length. Note that
the sheath and axis cylinder are of the same general intensity and so cannot
be distinguished. Specimen in distilled water ; birefringence positive in rela-
tion to nerve axis. Diameter of this fiber after stretching 10 p. Compen-
sator set for maximum brightness of nerve. Magnification 300 X.

Fig. 9. Same but with compensator plate rotated for extinction of nerve
retardation. Such brightening and extinguishing effects prove we are deal-
ing with birefringent properties.

Fig. 10. Another nerve from the same preparation. Diameter 12 p. Mag-
nification 300 x.

Fig. 11. Large group of nerve fibers from partly teased cockroach nerve
cord in saline containing 15% glycerine (2 hrs.). The form birefringence
of the protein is here masked sufficiently to give reversal of sign (metatropic
effect) ; the sign of birefringence here is then negative in relation to the
nerve axis. The many parallel light lines are due to the lipid components of
a number of nerve sheaths. Nerves of various diameters (3-8 p on plate
but somewhat shrunken by the glycerine medium). Magnification 500 X.

Fig. 12. A thoracic interganglionic connective of a cockroach in saline
with 15% glycerine (15 min.). The bright lines along the sides are pro-
duced by the neural lamella. The bright interior is produced by the super-
imposed effects of the lipid sheaths of many nerves. The neural lamella
is positive in relation to the nerve cord axis; the included bundle of nerves
is negative. Compensator set for maximum brightness. Magnification 45 X.

Fig. 13. Same but with compensator plate rotated for extinction of nerve
retardation. In this photomicrograph the positive birefringence of the
neural lamella is lost against the white background.

Fig. 14. Abdominal ganglion and interganglionic connectives of a mos-
quito larva. Fresh preparation in saline; birefringence positive in relation
to nerve axes. Compensator set for maximum brightness. Magnification
200 X.

Fig. 15. Same fresh preparation after 5 min. in saline containing 15%
glycerine. The sign has reversed and the nerve cord shrunken slightly.
Magnification 200 x.

Fig. 16. Portions of two interganglionic connectives of a cockroach after
18^ hours degeneration in saline containing 15% glycerine. Shows the break-
down of the sheath lipids into birefringent spheerites. The two bright lines
running diagonally through the center are produced by the neural lamellae
of the two connectives. The wavy bright line in the upper right quadrant
is produced by an air-filled trachea. Magnification 200 X.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate X)

Dec., 1944]

Milne: Behavior

311

NOTES ON THE BEHAVIOR OF BURYING BEETLES
(NICROPHORUS SPP.)

By Lorus J. Milne and Margery J. Milne
University of Pennsylvania and Beaver College

For several summers, ending with the season of 1928, the senior
author had observed species of Nicrophorus ( Coleoptera, Staphy-
linidae, Silphinae) burying small carcasses at Irondale (Halibur-
ton County), Ontario, Canada, in July. Some of his observa-
tions were published (Milne, 1928). Much more extensive and
detailed studies of burying beetles were made by Pukowski (1933)
in Europe and Leech (1935) in British Columbia. These authors
followed the life history from the time of burial, while the adults
cared for the young, through the three instars of larval life, the
prepupal and pupal periods, and made observations on emergence
and feeding of the adults. Difficulty was reported in observing
the burying behavior because of its nocturnal nature and because
Nicrophorus was not abundant enough to justify the risks of dis-
turbing beetles at work. Since the burying activities had been
watched repeatedly in daylight at Irondale, the present authors
made a return trip there in the summer of 1944, to prepare a
Kodachrome motion picture of the burying beetles in their work
above ground. Although the time available during the brief
vacation from war research greatly restricted the experimental
studies, it was possible to make further observations which extend
and clarify the behavior of these insects.

1. SPECIES INVOLVED AT IRONDALE

Six species of Nicrophorus have been collected at carrion in
this region by the authors, namely N. sayi Lap., N. orloicollis
Say, N. marginatus Fab., N. pustulatus Hersch., N. vespilloides
Hbst., and N. tomentosus Web. All these are easily distin-
guished in the field. Only tomentosus has the pronotum covered
with yellow, appressed hair (which often becomes abraded on the
two convexities of the disc). Only sayi and marginatus have
distinctly curved hind tibiae. Only vespilloides and tomentosus

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have the antennal clubs entirely black, the other species having
at least part of the club orange red ; these two species are also
considerably smaller in average size. Sayi and orbicollis are
very similar in markings, with the elytra mostly black, but with
orange red in a sub-basal, irregular crossbar and a subapical spot.
Marginatus, vespilloides and tomentosus are similar in markings,
the elytra being mostly orange red, the bands of color continuing
across both elytra, sometimes confluent also along the outer mar-
gins. Pustulatus is unique among these six species in having
the pronotum transversely oval, the others having a circular disc ;
it also has the sub-basal elytral spot small or absent, the subapical
bar often divided in two (sometimes lacking), the insect thus
having an even blacker facies than sayi or orbicollis. Of these,
only tomentosus and orbicollis * were numerous in late July of
1944, and on them most of the following notes are based.

2. METHODS OF STUDY

Irondale is a hilly region from which the original timber was
removed perhaps fifty years ago. Those areas which could be
freed of glacial erratics and which were reasonably level, have
been farmed (chiefly for grain). Other areas have been cleared
for pasture and kept available for sheep and cattle. Less level
land has been allowed to grow up again. The vegetation is typi-
cal Canadian zone, with hemlock, spruce, pine, paper birch, pop-
lar, spiraea, sweet. fern, sweet gale, mountain laurel, club mosses,
bracken, and an abundant lichen flora on exposed rocks and tree
trunks. Among the larger fauna are loons, whip-poor-wills, por-
cupines, groundhogs, skunks, red squirrels and chipmunks.
Most of the glacial lakes contribute to the Burnt River system,
deriving the name from the dark color of the water, due to leach-
ing of iron ore from rock substratum and to solution of decaying
coniferous and other xerophytic debris. Sphagnum is common
and pitcher plants and sundew reach large size.

To attract Nicrophorus, small dead animals were placed at
selected positions in a variety of ecological habitats. Each car-
cass was secured to a nearby stake by a two-foot length of fine steel
wire, to make recovery easy (c/. Milne, 1928). The animals used

* Referred to as N. pollinctor in Milne, 1928.

Dec., 1944]

Milne: Behavior

313

were chiefly deer mice (Peromyscus) , with some house mice
(Mus), shrews (Blarina), birds (robins, thrushes, sparrows,
wrens, domestic turkeys, etc., as found dead) and snakes. In
previous years larger carrion was tried, including groundhog
(Marmota) , skunk, dog, etc., but Nicrophorus was found to show
little interest in carcasses too large for them to bury. For spe-
cies observed at Irondale, a body the size of a robin is perhaps
the limit.

In open fields, such as pastures or where hay had been cut,
N. tomentosus and marginatus arrived to bury mice and snakes.
Competition with ants was frequent and the carcasses deterio-
rated considerably before the beetles could get them buried. In
leaf litter from birch and poplar, second growth woodland, N.
tomentosus and orbicollis were quick to bury mice and birds. In
coniferous duff the same species were somewhat slower in finding
carrion. No activity was observed on mice placed in low, wet
positions such as sedgy swales. Mice placed on particularly
hard ground or on bare rock, were transported by the beetles to
places where the soil was less packed and burial hence easier.
Desiccation of carcasses by the sun did not have any noticeable
effect on the interest in them shown by the beetles.

3. BEHAVIOK OF THE BEETLES

Nicrophorus exhibits a number of interesting behavior pat-
terns. The beetles fly to the general vicinity of the carrion, ap-
parently by smell (c/. Abbott, 1927a & 1927b; Milne, 1928).
Tomentosus is particularly accurate in locating the body before
alighting, buzzing through the bushes like a bumblebee. The
resemblance to Bombus is enhanced by the golden body hair, the
yellow inner surfaces of the elytra (which are held back to back
over the midline) and the creamy cast to the flying wings. All
Nicrophorus run about briskly, forcing their way through or
under tangled vegetation. If disturbed, all but tomentosus are
likely to either feign death, or run away a few feet to hide in
grass roots. Tomentosus takes to its wings and may alight in a
nearby bush, often standing on a slanting stem rather than the
broader surface of a leaf. Pukowski (1933) describes a similar
habit as part of the behavior of a lone Nicrophorus attracting a

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[Yol. lii

mate, the beetle climbing a stone or plant, elevating the abdomen
obliquely and extending it so much that the coriae are visible.
No indication of such was observed by the present authors, a lone
Nicrophorus regularly getting to work on the burial task and con-
tinuing so engaged until a mate arrived.

After a beetle has arrived at a dead animal, it characteristi-
cally examines the body with palpi and antennae and tests the size
of the carcass by trying to move it. The “carrying” action is
one of the most typical behavior patterns shown loy Nicrophorus.
Another activity closely related is that of “exploring” the sur-
rounding soil for a suitable spot in which the body may be in-
terred. A third procedure is to test the looseness of the soil by
“plowing” it. All of these behaviors are energetic, and are
shown by even a single Nicrophorus.

The carrying action of Nicrophorus demonstrates the strength
of the beetles and the vigor with which they proceed wtih their
task. To move a carcass, say forward in terms of a mouse’s body,
the beetle crawls under the head of the mouse, turns over on its
back, and lifts the mouse bodily over itself. The mouse inches
forward little by little, while the beetle slides slowly on its smooth
back under the mouse until it reaches the posterior end. There
it either emerges and runs around to the anterior end of the
mouse again, or turns over on its feet to crawl under the mouse,
the smooth dorsum of the beetle not disturbing the body. Such
effort is very strenuous, and a rest period or an exploring inter-
lude usually follows a few minutes of moving the carrion. A
single beetle rests more than when a mate has arrived. A slight
tug at the carcass usually stimulates it to renewed activity.
When two beetles are operating one or the other is usually carry-
ing the body while the mate explores or plows. No sexual differ-
ence in degree of activity could be noticed (c/. Wood, 1873;
Furneaux, 1893; Pukowski, 1933). Both members of a pair
were highly industrious under most circumstances. On level
ground devoid of major obstacles, a pair of beetles may transport
a full-grown mouse or shrew as rapidly as three or four feet per
hour, and keep up this pace for as much as two and a half hours,
the limit in every case observed being the distance necessary to
reach sufficiently soft ground for burial use.

Dec., 1944]

Milne: Behavior

315

The exploring behavior was very distressing to the observers.
After a shorter or longer period of work on a carcass, one or
both members of a pair will suddenly leave the body and run
away from it at the usual brisk pace. The beetles may go only
a few feet (seldom less), or as much as a few yards, and in some
cases ( tomentosus ) took to flight, only to return in a matter of
minutes. A specimen of N. tomentosus with a broken elytra! tip
was observed to fly off and back again four times within an hour.
Each time it flew out of sight. During the hour perhaps fifteen
minutes work was done in carrying the mouse and in plowing
in nearby soil, the remainder being either exploring the whole
surrounding area (perhaps fifteen feet in diameter), or resting
with head under a clump of grass, or absent altogether while
away on a flight. This procedure persisted even after the speci-
men was joined by agnate, continuing until the carrion had been
transported (mostly by the mate, in this instance), almost to the
area selected for burial use.

The plowing procedure loosens the earth. The beetle uses its
head as a plowshare or bulldozer, pressing into the earth perhaps
the depth of its stout body below the surface, then forcing its
way forward in an arc while maintaining its depth. The earth
is forced upward and crumbles. Any roots encountered are
either forced aside, or chewed through, but if numerous a new
burial site is sought out. Before a final area is considered satis-
factory, as many as twenty possible sites may be plowed and
found unsuitable. The final area may be many feet away from
the carrion, and the beetle (s) will alternately work on the car-
cass and run to the burial site to do another stretch of plowing.
The route taken between carrion and burial site is usually fairly
direct, and the body is carried along this line. How both mem-
bers of a pair can agree on a site was not determined, nor was
it at all clear how the beetles were able to keep the carcass moving
so uniformly (few exceptions) in one direction. The contrast
was very marked between the cooperation of Nicrophorus and the
great wastage of effort among ants trying to carry a large food
particle.

In a few instances, the beetle bait was placed on soft ground
and the Nicrophorus buried it where it lay. In all instances,

316 Journal New York Entomological Society [Vol. lii

however, the burial procedure was a localization of the plowing
action. The beetles plowed under the carrion, entering at one
side (or end) and emerging at the other, only to turn about and
complete the shuttling action. On each emergence at the side or
end of the carcass, a sizeable amount of earth was forced up, to
accumulate in a loose pile all around the body. Gradually the
earth from below the carrion was displaced to the side and the car-
cass settled into the ground. A continuation of the same activity
gradually drops the body below the surface of the ground, and the
procedure is continued until as much as two inches of earth are
over the carcass. The body is usually let into the ground at a
steep angle. Not only the earth under the carrion, but also that
within a few inches of it is plowed. Closer to the body the
ground is plowed to a greater depth. In its final site, a chamber
is cleared of earth below and to one side from the body. In the
chamber Nicrophorus remains after burial. The chamber may
be as much as two and a half inches long, an inch wide and deep,
and the walls are left packed in the original condition of the
earth, so that cave-ins are unlikely. Burial is usually completed
in five to eight hours, although the beetles will continue for days
if obstructions slow their work. Nicrophorus usually remain
with the carrion at least for many days (c/. Hatch, 1927b ; Milne,,
1928; Pukowski, 1933; Leech, 1935).

There seems to be a relationship between the size of the car-
rion, the size of the Nicrophorus species and the number of pairs
which can work in burying it. A full sized robin (in juvenile
plumage) was handled by a pair each of N. orbicollis and tomen-
tosus. A thirty-inch garter snake was buried by two pairs of
N. tomentosus and one of marginatus. In one instance a full-
grown deer mouse was interred by two pairs of A. tomentosus.
In all other instances only a single pair of a single species did
the work. Additional specimens of orbicollis arriving at a piece
of carrion where a pair were at work, usually crawled under the
body, only to emerge (promptly!) pursued by one of the original
pair. Several rough and tumble fights were witnessed in which
the newcomer resisted briefly the attack of the original 4 ‘ owner. ’ ’
Orbicollis coming to carrion where tomentosus was at work
usually drove the smaller species away. Tomentosus coming to

Dec., 1944]

Milne: Behavior

317

a carcass being carried or buried by orbicollis usually left with-
out any indication of expulsion. Tomentosus coming where
tomentosus were already at work sometimes helped awhile before
leaving, in only one instance (see above) remaining until the
body was completely interred. Thus tomentosus would seem to
make up in small size, greater numbers (see below) and more
sociable reactions for its inability to compete physically with the
larger species. Orbicollis becomes quite excited over expelling a
competitor and stridulates very audibly by rubbing the upper
surface of the abdomen against the under surface of the elytra
(cf. Morley, 1902). This same stridulation is observed when a
pair mate (3 to 4 seconds; cf. Pukowski, 1933), or when an ob-
struction continues to bar the way in moving a carcass. These
were the only occasions and the only species in which such noises
were noted. Pukowski (1933) indicates that pairs working on a
carcass keep in contact with each other by alternately emitting
“zirping” sounds.

Obstructions may be of a variety of types. Naturally encoun-
tered are situations where roots, stems, leaf petioles, etc., are suffi-
ciently anchored at both ends as to restrict an opening through
which the beetle is attempting to move the carrion. The beetle
discovering such an obstacle usually tries first to force it out of
the way, getting head and pronotum under the restraining strand,
feet in the vestiture of the carrion, and crawling ahead. Many
obstructions can be stretched sufficiently by this method to open
a suitable passage for further progress of the carcass. When
force proves inadequate, the beetle may settle down on the re-
straining piece and laboriously chew it through. Often periods
of chewing are alternated with episodes of drawing and pushing
at the carcass or strand, the chewing frequently having weakened
the obstruction sufficiently to let it give. When the obstacle,
proves too resistant to such treatment, soil is plowed from under
the carcass and the needed space obtained in this way.

Obstructions provided experimentally were usually very irri-
tating to the beetles but in no case did they leave permanently.
String tried to the leg or tail of a mouse was the commonest form
of restraint, and one which could be overcome by gnawing the
string until it broke under strain. Fine steel wire was an in-

318

Journal New York Entomological Society

[Vol. LII

superable difficulty. A carefully placed piece of rock, supported
partly on the ground and partly on the body of a mouse, proved
less of a problem. Although the rock weighed several pounds,
the pair of beetles working together were able to lift the rock
where it rested on the mouse sufficiently to push the carrion free
of its pressure and restraint.

One experimental situation gave a good demonstration of the
beetles’ behavior. A mouse laid out on fairly soft ground had a
string tied to one hind leg, the opposite end of the string being
fastened to a stake close to the mouse but in such a way that the
string was almost vertical and held the leg clear of the ground.
A pair of N. orbicollis proceeded to bury the mouse. They did a
fine job, the head of the mouse finally hanging almost vertically
downward into the hole the beetles had excavated. The hind leg
continued to be supported in its original position. The beetles
cleared away the earth until there was none below the head and
shoulders of the mouse for a distance equal to the thickness of a
beetle, and a space was also clear all around the mouse so that it
hung by its one leg over a sizeable, cup-shaped hole. With much
stridulation and rotation of the mouse around and around over
the hole, the beetles failed to go vertically upward to the support-
ing string. Every move in this direction ended in a trip out
the length of the mouse’s tail, to see that it was free. It was,
but the traffic became so heavy and the mouse so “ripe” that the
skin rolled off the tail distally like a glove finger. Finally the
beetles gnawed the tail off the mouse, severing it at the root.
Almost at once they found the supporting string, and after a
few abortive attempts to free the foot, one of the Nicrophorus
settled down to the task of gnawing it through. When the string
gave way, the mouse collapsed in a heap into the bottom of the
hole prepared for it, and burial was completed uneventfully
within half an hour.

Pukowski (1933) indicates that the beetles prepare a conical
hole below the carrion, always smaller than the body, and fold
the carcass as they draw it into the pit. The episode just de-
scribed would seem to indicate that Nicrophorus continue to
enlarge the hole for the carcass until it has been sunk to their
satisfaction, and that a conical hole and consequent folding may

Dec., 1944]

Milne: Behavior

319

be merely economy of effort. When carrion has considerable
length, as for example a snake, no folding was observed by the
present authors. The snake was let into the ground to a depth
of nearly two inches all along its length (except for the tail),
so that it was buried horizontally, in approximately the attitude
occupied previously on top of the earth. It is true that in the
case of snakes, burial advanced rapidly in the region from head
to anus, the tail being left out in the air for nearly twenty-four
hours after the remainder of the carcass was well under the sur-
face. The tail, with its smaller supply of food materials, was
obviously of much less interest to the beetles, forming chiefly an
obstruction to satisfactory burial. In several instances the last
few inches of a snake’s tail was chewed off, and the postanal
remainder pulled into the ground after putrefaction had rendered
it more plastic.

Heavy rain obstructed burying beetles much less than had been
expected. On several occasions Nicrophorus had begun to bury
bait placed on level, hard ground, when torrential downpours
drove the observers to nearby shelter. From the cabin the bait
could be seen almost or quite covered by water, sometimes to a
depth of an inch. No sign of the beetles was noted, yet within
fifteen minutes after the rain slackened enough for the ground
to drain off, the insects were busy in the wet earth, excavating,
plowing and tugging the carcass into their cavity.

The observers’ concern over the seeming desertion of the bait
whenever Nicrophorus went into an exploring episode, suggested
another experiment. As indicated above, the beetles frequently
explore a very sizeable area befor returning to the carcass. There
is no indication, however, that there is ever any difficulty in find-
ing the carcass again, since the beetles often return to it in an
almost straight line from a distance of a yard or two. On several
occasions, duplicate baits were tried, being mice of approximately
equal size and state of preservation, placed a foot apart on even
terrain. A Nicrophorus would come to one, test it for size, then
explore the surrounding ground for a suitable burial site. Dur-
ing the exploration the second mouse was usually discovered.
Almost without exception, the beetle examined the duplicate bait
without attempting to carry it, then hurried back to the mouse

320

Journal New York Entomological Society

[Vol. LII

previously discovered. In no case observed did the beetle (s)
desert the first bait in favor of the second. The same type of
experiment was tried on pairs which were busy burying a rela-
tively fresh mouse. A similar (or more odoriferous) mouse was
placed where they would surely find it during exploration trips.
In no case did the beetles desert the first-found carcass to more
than examine the second. It was quite obvious that the extra
carrion so nearby was a source of great distraction, but the recog-
nition of one body as distinct from another was most marked.

In only a few instances did Nicrophorus show any indication
of feeding on the carrion (cf. Furneaux, 1893 ; Lutz, 1921 ; Steele,
1927 ; Milne, 1928). Usually the beetles seemed to be in a hurry
to get the carcass interred. During daylight the need for rapid
burial was great, since blowflies came in considerable numbers,
laying living larvae if unmolested for a few minutes. The brisk
activities of the beetles and the frequent jerky movements of the
carcass have a deterrent effect on flies of some types (including
blowflies) but where obstacles prevented beetles from quickly get-
ting the carrion under ground during daylight hours, it was
obvious that little of the carcass would be available for other
than dipterous larvae. In many instances the beetles seemed to
realize this, and failed to complete burial. In some cases
Nicrophorus left fly-ridden carcasses sunk below the level of the
ground, covered by perhaps a quarter of an inch of loose earth.
A few days later such carrion was a squirming mass of fat fly
larvae. Rapidly buried bodies, on the other hand, are remark-
ably free of dipterous contamination. This may well be due to
the preference shown by beetles for operations in twilight, at
night or on cloudy (even rainy) days ( cf . Abbott, 1927b) or an
active eating of young maggots by the beetles (noted on a few
occasions; cf. Steele, 1927; Leech, 1935).

In one instance of Nicrophorus feeding, one member of a pair
took time off to investigate a small hole in the abdomen of a
“ripe” mouse, while the mate was busy excavating under the
carcass. For perhaps five minutes the feeding beetle worked
into the hole, until head and pronotum were inside the abdominal
wall. The viscera were explored rather superficially since the
position of the head could be discerned at all times by the moving

Dec., 1944]

Milne: Behavior

321

elevation it produced in the mouse’s skin. It was the observers’
opinion that the beetle was drinking rather than eating. Prior
to this feeding action, small flies had found the hole in the mouse ’s
abdominal wall a very interesting region, and considerable mois-
ture seemed to be present. After the beetle left the hole, there
was no liquid visible and no flies were attracted to the area.
During feeding, several blowflies ran against the posterior end
of the beetle, and were kicked away by violent movement of the
posterior legs. This kicking of molesting flies and ants seems a
common reaction in Nicrophorus.

Nicrophorus apparently discovers carrion entirely by smell,
while ants frequently locate freshly killed mice, seemingly as
part of routine foraging operations. Often ants had removed
the lips and nosetip of the rodent (the first part to be attacked
in all instances observed) before Nicrophorus arrived, but on
warm days (or nights) the beetles began to arrive within an hour
or less. Typical of the speed with which Nicrophorus gather
at a mouse is the following record made between six and ten
o ’clock one warm evening :

5 : 30 P.M.

Freshly killed mouse laid out in birch leaf litter.